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PLATE I

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AMERICAN INSECT

PLATE I.

SPHINX-MOTHS.

1 = Pholus pandoras.
2 = Smerinthus geminatus.

3=Ampelophaga versicolor.

4=Marumba modesta.

5
= Hemaris thysbe.
6=Thyrcus abbotti.
.

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 American jjiature
Group I. Classification of Nature

AMERICAN INSECTS

VERNON L. KELLOGG
Pro/esior of Entomology and Lecturer on Bionomics
in Leland Stanford Jr. University

WITH MANY ORIGINAL ILLUSTRATIONS

BY

MARY WELLMAN

SECOND EDITION, REVISED

NEW YORK
HENRY HOLT AND COMPANY
1908
 Copyright, 1904. 1908,
BY
HENRY HOLT AND COMPANY

ROBERT DRUMMOND COMPANY, PRINTERS. KEW YORK

 TO

JOHN HENRY COMSTOCK

 PREFATORY NOTE
TO SECOND EDITION, REVISED
IN this new and revised edition of American Insects a detailed analytical
table of contents has been substituted for the simple list of chapter titles
used in the first edition,and an additional chapter (Chapter XIX) on the
subject of insect behavior and psychology has been added. While descrip-
tive accounts of the reflexes and instincts of insects are to be found on
almost every page of the book insect instinct is indeed one of the primary

subjects of the book the author has believed that a special discussion and
attempt at analysis of the springs and control of insect behavior would be
of interest to the reader. This special though necessarily all too condensed
and brief treatment of the subject has therefore been introduced into the
present edition. V. L. K.

STANFORD UNIVERSITY,
March 26, 1908.
 PREFATORY NOTE
IF man were not the dominant animal in the world, this would be the
Age of Insects. Outnumbering in kinds the members of all other groups
of animals combined, and showing a wealth of individuals and a degree
of prolificness excelled only by the fishes among larger animals, and among
smaller animals by the Protozoa, the insects have an indisputable claim on
the attention of students of natural history by sheer force of numbers. But
their claim to our interest rests on securer ground. Their immediate and
important relation to man as enemies of his crops, and, as we have come to
know only to-day, as it were, as a grim menace to his own health and life
this capacity of insects to destroy annually hundreds of millions of dollars'
worth of grains and fruitsand vegetables, and to be solely responsible for
the dissemination of some of the most serious diseases that make man to
suffer and die, forces our attention whether we will or not. Finally, the
amazing variety and specialization of habit and appearance, the extraor-
dinary adaptations and "shifts for a living" which insects show, make a
claim on the attention of all who harbor the smallest trace of that "scientific

curiosity" which leads men to observe and ponder the ways and seeming of
Nature. Some of the most attractive and important problems which modern
biological study is attacking, such as the significance of color and pattern,
the reality of mechanism and automatism in the action and behavior of
animals as contrasted with intelligent and discriminating performances,
the statistical and experimental study of variation and heredity, and other sub-

jects of present-day biological investigation,

are finding their most available
material and data among the insects.
This book is written in the endeavor to foster an interest in insect biology
on the part of students of natural history, of nature observers, and of general
readers; it provides in a single volume a general systematic account of all
the principal groups of insects as they occur in America, together with special
accounts of the structure, physiology, development and metamorphoses', and
of certain particularly interesting and important ecological relations of insects
with the world around them. Systematic entomology, economic entomology,
and what may be called the bionomics of insects are the special subjects of
the matter and illustration of the book. An effort has been made to put
the matter at the easy command of the average intelligent reader; but it has
been felt that a little demand on his attention will accomplish the result
more satisfactorily than could be done with that utter freedom from effort
 Prefatory Note

with which some Nature-books try to disseminate knowledge. The few

technical terms used are all explained in the text in connection with their
first use, and besides are inserted in the Index with a specific reference, in
black-faced type, to the explanation. So that the tyro reading casually in
the book and meeting any of these terms apart from their explanation ha?
only to refer to the Index for assistance. Readers more interested in account?
of the habits and kinds of insects than in their structure and physiology
will be inclined to skip the first three chapters, and may do so and still find
book "easy reading" and, it is hoped, not devoid
the rest of the of entertain-
ment and advantage. But the reader is earnestly advised not to spare the
attention especially needed for understanding these first chapters, and
little

thus to ensure for his later reading some of that quality which is among
the most valued possessions of the best minds.
In preparing such a book as this an author is under a host of obligations

to previous writers and students which must perforce go unacknowledged.

Some formal recognition, however, for aid and courtesies directly tendered
by J. H. Comstock of Cornell University, whose entomological text-books
have been for years the chief sources of knowledge of the insects of this
country, I am able and glad to make. To my artist, Miss Mary Wellman,
for her constant interest in a work that must often have been laborious and

wearying, and for her persistently faithful endeavor toward accuracy, I extend
sincere thanks. To Mrs. David Starr Jordan, who read all of the manuscript
as a "general reader" critic, and to President Jordan for numerous sugges-
tions I am particularly indebted. For special courtesies in the matter of
illustrations (permission to have electrotypes made from original blocks)
I am obliged to Prof. F. L. Washburn, State Entomologist of Minnesota (for

nearly one hundred and fifty figures), Prof. M. V. Slingerland of Cornell

University, Dr. E. P. Felt, State Entomologist of New York, Mr. Wm.
Beutenmuller, editor of the Journal of, the New York Entomological Society,
and Dr. Henry Skinner, editor of the Entomological News.
VERNON L. KELLOGG.
STANFORD UNIVERSITY, CALIFORNIA,
June i, 1904.
 CONTENTS
CHAPTER I
PAGE
STRUCTURE AND SPECIAL PHYSIOLOGY OF INSECTS i
Structural characteristics of the class Insecta, 3. External anatomy, 4.

Body-wall, 4. Divisions of body, 5. Mouth-parts, 8. Wings, 9. Size and

form of body, 12 Internal anatomy, 13. Muscles, 13. Alimentary canal,
13. Reproductive system, 14. Circulatory system, 16. Respiratory system,
19. Nervous system, 20. Special sense-organs, 24. Insect psychology, 33.

CHAPTER II

DEVELOPMENT AND METAMORPHOSIS 35

Eggs and embryonic development, 36. Post-embryonic development, 40.
Development without metamorphosis, 41. Development with incomplete meta-
morphosis, 41. Development with complete metamorphosis, 43. Internal
changes during development, 47. Significance of development, 49.

CHAPTER III

CLASSIFICATION OF INSECTS 52
Various schemes of classification into orders, 52. Analytical key to the orders
of insects, 54.

CHAPTER IV

THE SIMPLEST INSECTS (ORDER APTERA) 58

Fish-moths and springtails, 58. Primitiveness among insects, 59. Struc-
tural characteristics, 59. Key to the suborders, 60. Thysanura, 60. Key to
the families of Thysanura, 60. Key to genera of Lepismidae, 61. Collembola,
62. Key to families of Collembola, 63.

CHAPTER V
MAY-FLIES (ORDER EPHEMERIDA) AND STONE-FLIES (ORDER PLECOPTERA) 65
May-fly swarms, 65. Structure of adults, 68.
Life-history, 66. Stone-flies,
70. Life-history, 71. Structure of adults, 71. Table of North American
genera of Plecoptera, 73.
 Contents

CHAPTER VI
PAGE
DRAGON-FLIES AND DAMSEL-FLIES (ORDER ODONATA) 75
Characteristics and distribution of dragon-flies, 76. Structure of adults, 79.
Habits, 81. Life-history, 84. Methods of collecting and studying, 87. Various
kinds of dragon-flies, 89. Keys to suborders, 89. Key to families of Zygoptera,

89. Key to families of Anisoptera, 91.

CHAPTER VII

TERMITES OR WHITE ANTS (ORDER ISOPTERA) 99

Characteristics and structure of Termites, 99. Life-history, 101. Key to

genera, 102. Habits and characteristics of various kinds of termites, 102.

Termites of Africa, 106. The problem of caste determination, 108.' The
Embiidae, 109.

CHAPTER VIII

BOOK-LICE AND BARK-LICE (ORDER CORRODENTIA) AND THE BITING BIRD-LICE

(ORDER MALLOPHAGA)
Structure and life-history of book-lice, in.
m
Keys to families and genera,
112. Characteristics and life-history of bird-lice, 113. Problems of distri-
bution, 1 1 6.
Keys- to suborders, families, and genera, 118. Various species,
119.

CHAPTER IX
THE COCKROACHES, CRICKETS, LOCUSTS, GRASSHOPPERS, AND KATYDIDS (ORDER
ORTHOPTERA) 123
Sounds of crickets, etc., 123. Key to families, 126. Cockroaches or Blat-
tidae, 126. Mantidae, 129. Phasmidae, 132. Key to genera of Phasmidas,
132. Locusts (Acridiidae), 133. Sounds of locusts, 134. Life-history of
locusts, 136. Key to subfamilies of Acridiidae, 136. Rocky Mountain locust,
137. Various kinds of locusts, 140. Locustidae, 149. Various kinds, 150.
Crickets (Gryllidae), 157. Sound-making of crickets, 157. Ear-wigs (Forficu-
lidae), 162.

CHAPTER X
THE TRUE BUGS, CICADAS, APHIDS, SCALE-INSECTS, ETC. (ORDER HEMIPTERA),
AND THE THRIPS (ORDER THYSANOPTERA) ^3
Characteristics of Hemiptera, 164. Key to suborders, 165. Key to families
of Homoptera, 166. Cicadas (Cicadidae), 166. Tree-hoppers (Membracidas)
and lantern-flies (Fulgoridae), 168. Leaf-hoppers (Jassidae), 169. Spittle
insects (Cercopidae), 170. Jumping plant-lice (Psyllidae), 171. Plant-lice
(Aphidiidae), 171. Grape-phylloxera, 176. Scale-insects (Coccidas), 180. San
Jose scale, 181. Remedies for scale-insects, 189. Mealy-winged flies (Aleyro-

didae), 190. Key to families of Heteroptera, 194. Water-striders (Hydrota-

tidae), 196. Water-boatmen (Corisidae), 198. Back-swimmers (Notonectidac),
 Contents xi

PAGE
198. Water-creepers (Naucoridae), 199. Giant water-bugs (Belostomatidae),
199. Water-scorpions (Nepidae), 201. Toad-bugs (Galgulidae), 202. Shore-
bugs (Saldidae), 202. Assassin-bugs (Reduviidae), 203. Thread-legged bugs
(Emesidae), 204. Damsel-bugs (Nabidae), 204. Bedbugs (Acanthiidae), 205.
Key to families of plant-feeding Heteroptera, 207. Lace-bugs (Tingitidae), 207.
Flat-bugs (Aradidae), 208. Flower-bugs (Capsidae), 209. Red-bugs (Pyrrho-
coridae), Chinch-bugs and others (Lygaeidae), 211. Squash-bugs and
210.
others (Coreidae), 213. Stilt-bugs (Berytidae), 214. Shield-bodied bugs (Pen-
tatomidae), 214. Lice (Pediculidae), 216. Thrips (Thysanoptera), 219.

CHAPTER XI
THE NERVE-WINGED INSECTS (ORDER NEUROPTERA), SCORPION-FLIES (ORDER
MECOPTERA), AND CADDIS-FLIES (ORDER TRICHOPTERA) 223
Key to the families of Neuroptera, 224. Key to the genera of Sialidae, 224.

Lace-winged flies (Chrysopidae), 228. Aphis-lions (Hemerobiidae), 229. Ant-

lions (Myrmeleonidas), 230. Key to subfamilies, 231. Key to genera of Myr-
meleoninae, 232. Key to Ascalaphinae, 233. Snake-flies (Raphidiidae), 233.

Mantispidae, 234. Coniopterygidae, 235. Scorpion-flies and others (Mecoptera),

235. Key to genera, 236. Caddis-flies (Trichoptera), 239. Cases of caddis-
flies, 240. Life-history, 241. Key to families (adults), 244. Key to families
(larvae), 244.

CHAPTER XII
THE BEETLES (ORDER COLEOPTERA) 246
External structure, 247. Internal structure, 248. Character of antennae and
legs, 250. Key to sections and tribes, 251. Key to families of Adephaga, 252.
Tiger-beetles (Cicindelidae), 252. Predaceous ground-beetles (Carabidae), 253.
Diving beetles (Dyticidae), 255. Whirligig beetles (Gyrinidae), 257. Key to
families of Clavicornia, 258. Water-scavenger beetles (Hydrophilidae), 258.
Rove-beetles (Staphylinidae), 260. Carrion-beetles (Silphidae), 261. Grain-
beetles and others (Cucujidae), 262. Larder-beetles and others (Dermestidae),
263. Water-pennies (Parnidae), 264. Beaver-beetles (Platypsyllidae), 265. Key
tofamiliesof Serricornia, 265. Metallic wood-borers (Buprestidae), 265. Click-
beetles (Elateridae), 267. Fire-flies (Lampyridae), 269. Checker-beetles (Cleridae),

270. Drug-store beetles and others (Ptinidae), 271. Key to families of Lamellicor-
nia, 272. Stag-beetles (Lucanidae), 272. Leaf-chafers and others (Scarabaei-
dae), 273. families of Tetramera, 277.
Key to Leaf-eating beetles (Chrysome-
lidae), 277. Pea- and bean-weevils (Bruchidae), 281. Long-horn boring beetles
(Cerambycidae), 282. Lady-bird beetles (Coccinellidae), 286. Key to families
of Heteromera, 288. Darkling ground-beetles (Tenebrionidae), 288. Blister-
and Wasp-beetles (Stylopidae), 293. Key to fami-
oil-beetles (Meloidae), 289.
lies of Rhynchophora, 294. Scarred snout-beetles (Otiorhynchidae), 295. Cur-
culios and weevils (Curculionidae), 295. Rice- and grain-weevils (Calandridae),
297. Engraver beetles (Scolytidae), 298.

CHAPTER XIII
TWO-WINGED FLIES (ORDER DIPTERA) 301
Characteristics of the Diptera, 301. Table to suborders and sections, 303.
Key to families of Nematocera, 304. Mosquitoes (Culicidae), 305. Mosquitoes
xii Contents

PAGE
and human disease, 308. Midges (Chironomidae), 310. Black-flies (Simuliidae),

313. Net-winged midges (Blepharoceridae), 314. Dixidae, 318. Moth-flies

(Psychodidae), 319. Crane-flies (Tipulidae), 321. Gall-midges (Cecidomyidae),

322. Fungus-flies (Mycetophilidae), 324. March-flies (Bibionidae), 325.
Orphnephilidas and Rhyphidae, 327. Section Brachycera, 327. Key to

families, 327. Horse-flies (Tabanidae), 328. Soldier-flies(Stratiomyidae),

329. Snipe-flies (Leptidae), 330. Midas flies (Midaidae), 330. Robber-flies
(Asilidae), 330. Key to families of Brachycera, 332. Bee-flies (Bombyliidae),

333. Dance-flies (Empididae)j 334. Long-legged flies

(Dolichopodidae), 335.
Wasp-flies (Conopidae), 336. Bot-flies (Oestridae), 337. Flower-flies (Syrphi-

dae), 339. Calyptrate Muscidae, 341. Key to subfamilies, 341. House-flies,

etc., 342. Tachina flies, 345. Acalyptrate Muscidae, 346. Ephydridse, Pio-
philidae, Drosophilidae, Trypetidae, Oscinidae, etc., 347. Suborder Pupipara,
351. Key to families, 351. Sheep-ticks, bat-ticks, bee-lice, etc., 351. Order
- to families, 355.
Siphonaptera, 353. Fleas, 353. Key

CHAPTER XIV
MOTHS AND BUTTERFLIES (ORDER LEPIDOPTERA) 358
Structural characteristics, 358. Life-history, 360. Classification into sub-
orders, 364. Key to superfamilies and families of moths, 367. Jugate moths
(Micropterygidae), 371. Ghost-moths (Hepialidae), 372. Microlepidoptera, 374.
Clothes-moths (Tineidas), 374. Pryalidina, 376. Plume-moths and others
(Pterophoridae), 377. Close-wings (Crambidae), 377. Meal-moths, flour-moths,
bee-moths, and others (Pyralidae), 378. Leaf-rollers (Tortricidae), 379. Flannel-
moths (Megalopygidae), 383. Slug-caterpillar moths (Eucleidae), 384. Car-
penter-moths (Cossidae), 385. Bag-worm moths (Psychidae), 385. Smoky-
moths (Pyromorphidae), 386. Clear-wing moths (Sesiidae), 388. Puss-moths,
handmaid-moths, prominents, etc. (Notodontidae), 392. Inchworm-moths
(Geometrina), 395. Owlet-moths (Noctuidae), 399. Tussock-moths (Lyman-
triidae), 404. Oak-moths (Dioptidas), 407. Pericopidae, 407. Wood-nymph
moths (Agaristidae), 407. Footman-moths (Lithosiidae), 409. Zygaenid moths.
(Syntomidae), 410. Tiger-moths (Arctiidas), 411. Tent-caterpillar moths
(Lasiocampidae), 415. Bombyx moths (Saturniina), 417. Silkworm-moths,
418. Mulberry silkworm, 429. Sphinx-moths (Sphingidae), 431. Butterflies,
439. Key to families of butterflies, 441. Giant-skippers (Megathymidae), 441..
Skipper-butterflies (Hesperidae), 442. Blues, coppers, and hair-streaks (Lycae-
nidae), 443. Cabbage-butterflies and others (Pieridse), 444. Swallow-tails
(Papilionidae), 446. Brush-footed butterflies (Nymphalidae), 450.

CHAPTER XV
SAW-FLIES, GALL-FLIES, ICHNEUMONS, WASPS, BEES, AND ANTS (ORDER HYMEN-
OPTERA) 459.
Structural characteristics, 459. Life-history, 461. Key to superfamilies and
families, 463. Saw-flies and slugs (Tenthredinidae), 464. Horntails (Siricidae),
466. Gall-flies (Cynipidae), 467. Parasitic Hymenoptera (Proctotrypoidae,
Chalcidiidae, Ichneumonidae), 477. Fig-insects, 487. Wasps, solitary and social,
490. Classification into superfamilies and families, 490. Habits and instincts
 Contents xiii

PAGE
of solitary wasps, 491. Velvet-ants (Mutillidse), 497. Cuckoo-flies (Chrysididae),
498. Mason- or potter-wasps, 498. Eumenidse, 498. Digger-wasps (Sphecidae,
Larridas, Bembecidas, Pompilidae), 499. Wood-mining wasps (Mimesidae, Pem-
phredinidae, Crabronidae), etc., 502. Social wasps (Vespidas), 503. Key to

genera, 503. Life-history of community of yellow-jackets, 503. Bees, 510.

Characteristics, 511. Solitary bees, 513. Mining-bees and carpenter-bees, 513.
Mason-bees and potter-bees and leaf-cutters, 514. Mining-bees, 516. Social
bees, 517. Bumblebees, 517. Honey-bees, 520. Life-history of community, 521.
Ants (Formicina), 533. Characteristics and life-history, 535. Key to families,
540. Poneridae, 540. Myrmicidae, 541. Camponotidae, 545. Artificial nests,

548. Myrmecophily, 552. Problems of ant behavior, 554.

CHAPTER XVI
INSECTS AND FLOWERS 562
Relations between plants and insects, 562. Cross-pollination in flowers, 563.
Means of avoiding self-fertilization, 565. Specialization for cross-pollination,
566. Uses of nectar and odor, 567. Modifications of insect visitors, 569. Par-
ticular cases of flower specialization for cross-pollination, 571. Tubular corollas,
571. Irregular tubular flowers, 572. Cross-pollination in Asclepias, 573.
Cross-pollination of Araceae and Aristolochiaceae, 575. Cross-pollination of
orchids, 575. Cross-pollination of Yucca by Pronuba, 576. Origin of speciali-
zations for cross-pollination, 579.

CHAPTER XVII

COLOR AND PATTERN AND THEIR USES 583

Wide distribution of color and pattern among insects, 583. Explanations of
some color phenomena in insects, 583. How color in organisms is produced,
586. Classification of insect colors, 587. Color patterns of the butterflies and
moths produced by scales, 589. Characteristics of the scales, 589. Ontogenetic
appearance of color pattern in insects, 596. General protective resemblance, 599.
Variable protective resemblance, 599. protective resemblance, 602.
Special
Warning colors, 604. Terrifying appearances, 605. Directive coloration, 607.
Mimicry, 608. Criticisms of hypotheses of color use, 611.

CHAPTER XVIII

INSECTS AND DISEASE 615

Economic between insects and man, 615. Dissemination of human
relations
diseases by insects, 616. Mosquitoes and malaria, 617. Mosquitoes and yel-
low fever, 630. Mosquitoes and filariasis, 632.

CHAPTER XIX
REFLEXES, INSTINCTS, AND INTELLIGENCE 635
Theories of insect behavior, 635. Points of view of Loeb and Jennings; tro-

pisms and method of trial and error, 635. Distinguishing among reflexes,
xiv Contents

PAGE
instincts, and intelligence, 636. Reflexes and tropisms, 638. Davenport's
analysis of behavior of Poduridae, 639. The swarming reflex of honey-bees, 639.
Reflexes of silkworm-moths, 640. Instincts, 641. Complex behavior of solitary
wasp, 643. Fabre's experiments and conclusions, 643. Peckham's experiments
and conclusions, 650. An increasing mass of evidence favoring mechanical
explanation of insect behavior, 655.

APPENDIX
COLLECTING AND REARING INSECTS 656
Collecting equipment, 656. When and how to collect, 660. Rearing insects,
661. Aquarium, 665.

INDEX 669
 AMERICAN INSECTS
CHAPTER I

THE STRUCTURE AND SPECIAL

PHYSIOLOGY OF INSECTS
ERHAPS no more uninteresting matter, for
the general reader or entomological amateur,
can be written about insects than a descrip-
tive catalogue of the parts and pieces of the
insect body. And such matter is practically
useless because it doesn't stick in the reader's
mind. If it is worth while knowing the
intimate make-up of a house-fly's animated little body, it is worth
getting this knowledge in the only way that will make it real, that is,
by patient and eye-straining work with dissecting-needles and micro-
scope. This book, anyway, is to try to convey some information about
the kinds and ways of insects, and to stimulate interest in insect life, rather
than to be a treatise on insect organs and their particular functions. Life
is, to be sure, only the sum of the organic functions, but this sum or com-

bination has an interest disproportionate to that of any of its component

parts, and has an aspect and character which cannot be foretold in any com-
pleteness from ever so careful a disjoined study of the particular functions.
And so with the body, the sum of the organs: it is the manner and seeming
of the body as a whole, its symmetry and exquisite adaptation to the special
habit of life, the fine delicacy of its colors and pattern, or, at the other
extreme, their amazing contrasts and bizarrerie, on which depend our first
interest in the insect body. A second interest, although to the collector and
amateur perhaps the dominant one, comes from that recognition of the
differences and resemblances among the various insects which is simply
the appreciation of kinds, i.e., of species. This interest expanded by oppor-
tunity and observation and controlled by reason and the habit of order and
arrangement is, when extreme, that ardent and much misunderstood and
scoffedat but ever-impelling mainspring of the collector and classifier.
2 The Structure and Special Physiology of Insects

Of all entomologists, students of insects, the very large majority are col-
lectors and classifiers, and of amateurs apart from the few who have "crawl-
"
eries" and aquaria for keeping alive and rearing worms " and water-bugs
and the few bee-keepers who are more interested in bees than honey, prac-
tically all are collectors and arrangers. So, as collecting depends on a
knowledge of the life of the insect as a whole, and classifying (apart from
certain primary distinctions) on only the external structural character of
the body, any detailed disquisition on the intimate character of the insec-
tean insides would certainly not be welcome to most of the users of this
book.
That insects agree among themselves in some important characteristics
and differ from all other animals in the possession of these characteristics
is implied in the segregation of insects into a single great class of animals-
Class here is used with the technical meaning of the systematic zoologist-
He says that the animal kingdom is separable into, or, better, is composed
of several primary groups of animals, the members of each group possessing
in common certain important and fundamental characteristics of structure
and function which are lacking, at any rate in similar combination, in all

other animals. These primary groups are called phyla or branches. All
the minute one-celled animals, for example, compose the phylum Protozoa

(the simplest animals); all the starfishes, sea-urchins, sea-cucumbers, and

which have the body built on a radiate plan and have no back-
feather-stars,
bone, and have and do not have certain various other important things,
compose the phylum or branch Echinodermata all the back-boned ani-
;

mals and some few others with a cartilaginous rod instead of a bony column
along the back compose the class Chordata; all the animals which have
the body composed of a series of successive rings or segments, and have

pairs of jointed appendages used as feet, mouth-parts, feelers, etc., aris-

ing from these segments, compose the phylum Arthropoda. There are
still other phyla but I am not writing a zoology. The insects are Arthro-
poda; and any one may readily see it is most plainly seen in such forms as
a locust, or dragon-fly, or butterfly, and less plainly in the concentrated
knobby little body of a house-fly or bee that an insect's body shows the
characteristic arthropod structure; it is made up of rings or segments, and
the appendages, legs for easiest example, are jointed. An earthworm's
body is made up of rings, but it has no jointed appendages. A worm is

therefore not an arthropod. A crayfish, however, is made up of distinct

successive body-rings, and itsand other appendages are jointed. And
legs
so with crabs and lobsters and shrimps. And the same is true of thousand-
legged worms and centipeds and scorpions and spiders. All these creatures,
then, are Arthropods. But they are not insects. So all the back-boned
animals, fishes, amphibians, reptiles, birds, and mammals are Chordates,
 The Structure and Special Physiology of Insects 3

but they are not all birds. The phylum Chordata is subdivided into or
composed of the various classes Pisces (fishes), Aves (birds), etc. And
similarly the phylum Arthropoda is composed of several distinct classes,
the Crustacea, including the crayfishes, crabs, shrimps, lobsters,
water-fleas, and barnacles;the Onychophora, containing a single genus

(Peripatus) of worm-like creatures; the Myriapoda, including the thousand-

legged worms and centipeds; the Arachnida, including the scorpions, spiders,
mites, and ticks; and finally the class Insecta (or Hexapoda, as it is some-
times called), whose members are distinguished from the other Arthro-

antennse

ovipositor

femur*
tibia

tarsal segments

FIG. i. Locust (enlarged) with external .parts named.

pods by having the body-rings or segments grouped into three regions, called
head, thorax, and abdomen, by having jointed appendages only on the body-
rings composing the head and thorax (one or two pairs of appendages may
occur on the terminal segments of the abdomen) and by breathing by means
,

of air-tubes (tracheae) which ramify the whole interior of the body and

open on its surface through paired openings (spiracles). The insects also
have three pairs of legs, never more, and less only in cases of degeneration,
and by this obvious character can be readily distinguished from the Myria-
pods, which have many pairs, and the Arachnids, which have four pairs.
Centipeds are not insects, nor are spiders and mites and ticks. What
are insects most of this book is given to showing.
To proceed to the classifying of insects into orders and families and
genera and species inside of the all-including class is the next work of the
collector and classifier. And for this if for no other reason some further
knowledge of insect structure is indispensable. The classification rests
4 The Structure and Special Physiology of Insects

mostly on resemblances and differences in corresponding parts of the body,

apparent in the various insect kinds. What these parts are, with their names
and general characters, and what their particular use and significance are,
may be got partly from the following brief general account, and partly from
the special accounts given in connection with special groups of insects else-
where in this book. A
little patience and concentration of attention in

the reading of the next few pages will make the reader's attention to the
rest of the book much simpler, and his understanding of it much more
effective.
The outer layer of the skin or body-wall of an insect is called the cuticle,
and in most insects the cuticle of most of the body is firm and horny in char-

to show chitin-
FIG. 2. Longitudinal section of anterior half of an insect, Menopon titan,
ized exoskeleton, with muscles attached to the inner surface. (Much enlarged.)

acter,due to the deposition in it, by the cells of the skin, of a substance called
chitin. This firm external chitinized * cuticle (Fig. 2) forms an enclosing
exoskeleton which serves at once to protect the inner soft parts from injury

p IG git of body
' -wall, greatly magnified,
of larva of blow -fly, Calliphora erythrocephala,
^
to show attachment of muscles to inner surface.

and to afford rigid points of attachment (Figs. 2, 3 and 4) for the many small
but strong muscles which compose the insect's complex muscular system.
Insects have no internal skeleton, although in many cases small processes
in the thorax or part
project internally from the exoskeleton, particularly
* It is not certainly known whether the cuticle is wholly secreted by the skin cells, or

is in part composed of the modified external ends of the cells themselves.

 The Structure and Special Physiology of Insects 5;

of the body bearing the wings and legs. Where the cuticle is not strongly
chitinized it is flexible (Fig. 6), thus
permitting
the necessary movement or play of the rings
of the body, the segments of the legs, antennae
and mouth-parts, and other parts. The small
portions of chitinized cuticle thus isolated or
made separate by the thin interspaces or sutures

w.

FIG. 5.

FIG. 4. Diagram of cross-section through the thorax of an insect to show leg and wing
muscles and their attachment to body-wall, h., heart; al.c., alimentary canal; v.n.c.
ventral nerve-cord; w., wing; /., leg; w., muscles. (Much enlarged after Graber.)
;

FIG. 5. Left middle leg of cockroach with exoskeleton partly removed, snowing muscles.
(Much enlarged; after Miall and Denny.)

are called sclerites, and many of them have received specific names, while
their varying shape and character are made use of in distinguishing and
classifying insects.

FIG. 6. Chitinized cuticle from dorsal wall of two body segments of an insect, showing
sutures (the bent places) between segmental sclerites. Note that the cuticle is not
less thick in the sutures than in the sclerites, but is less strongly chitinized (indi-
cated by its paler color).

The whole body is composed fundamentally of successive segments

(Figs, i which may be pretty distinct and similar, as in a caterpillar
and 7),
or termite or locust, or fused together, and strongly modified, and hence
dissimilar, as in a house-fly or honey-bee. The segments, originally five
or six, in all insects wholly fused to form a
composing the head, are single
box-like cranium, while the three segments which compose the thorax are
in most forms so fused and modified as to be only with difficulty distinguished
as originally independent body-rings. On the other hand, in most insects
6 The Structure and Special Physiology of Insects

the segments of the abdomen retain their independence and are more or

compound eye,

antennae^
'

prothorax^

labial

palpi
proboscis'
\ \metathoraz
* \ mesothorax

tarsal segments

FlG. 7. Body of the monarch butterfly, Anosia plexippus, with scales removed to show
external parts. (Much enlarged.)

less similar, thus preserving a generalized or ancestral condition. On the

head are usually four pairs of jointed appendages (Fig. 8), viz., the
antennae and three pairs of mouth-parts,
known as mandibles, maxillae, and labium or
.aur/t, under-lip. Of these the mandibles in most
cases are only one-segmented, while the two
members of the labial pair have fused along
their inner edges to form the single lip-like
labium. The so-called upper lip or labrum,
closing the mouth above, is simply a fold of
the skin, and is not homologous, as a true
appendage or pair of appendages, with the
other mouth-parts. In some insects with highly
modified mouth structure certain of the parts

FIG. 8.
7" ofr head
Dorsal aspect
,
may* be wholly
*,
lost, as is true of the mandibles
of dobson-fly, Corydalis cor- m the case of all the butterflies. The head
nuta, female, showing mouth- he
k ears ai so arge compound eves and the
t i
parts. lb., labrum, removed; ,, . . ... ...

md., mandible; mx., maxilla;

smaller simple eyes or ocelli (for an account of
U., labium; gl, glossae of la- the eyes see p. 30). Attached to the thorax are
bium; st., stipes of maxilla; ., ri.
i_-v i j j .*.

threg pairs of legs, which are jointed appendages,

nixp., palpus of maxilla; ant.,
antenna. homologous in origin and fundamental struc-
ture with the mouth-parts and antennae, and two pairs of wings (one or
 The Structure and Special Physiology of Insects 7

both pairs wanting) which are expansions of the dorso-lateral

may be
skin body-wall, and are not homologous with the jointed ventral
or

appendages. The thorax usually has its first or most anterior segment,
the prothorax, distinct from the other two and freely movable, while
the hinder two, called meso- and meta-thoracic segments, are usually
enlarged and firmly fused to form a box for holding and giving attachment
to the numerous strong muscles which move the wings and legs. The
abdomen usually includes ten or eleven segments without appendages or
projecting processes except in the case of the last two or three, which bear
in the female the parts composing the egg-laying organ or ovipositor, or

FIG. 9. FIG. 10.

FIG. Head, much enlarged, of mosquito, Culex sp., showing piercing and sucking
9.
mouth-parts. (After Jordan and Kellogg.)
FIG. 10. Head and mouth-parts of honey-bee, much enlarged. Note the short, trowel-
like mandibles for moulding wax when building comb, and the extended proboscis
for sucking flower-nectar. (Much enlarged.)

in certain insects the sting, and in the male the parts called claspers, cerci,
etc., which are used in mating. the On abdomen
are usually specially notice-
able, as minute paired openings on the lateral aspects of the segments, the

breathing-pores or spiracles, which admit air into the elaborate system of

tracheae or air- tubes, which ramify the whole internal body (see p. 19).
Of all these external parts two groups are particularly used in schemes
of classification because of their structural and physiological importance
in connection with the special habits and functions of insect life, and because
8 The Structure and Special Physiology of Insects

of the pronounced modifications and differences in their condition: these

are the mouth-parts and the wings.
Insects exhibit an amazing variety in food-habit: the female mosquito likes
blood, the honey-bee and butterfly drink flower-nectar, the chinch-bug sucks
the sap from corn-leaves, the elm-leaf beetle and maple -worm bite and chew
I he leaves of our finest shade-trees, the carrion-beetles devour decaying
animal matter, the house-fly laps up sirup or rasps off and dissolves loaf-
sugar, the nut- and grain-weevils nibble the
dry starchy food of these seeds, while the
apple-tree borer and timber-beetles find
sustenance in the dry wood of the tree-
trunks. The biting bird-lice are content
with bits of hair and feathers, the clothes-
moths and carpet-beetles feast on our rugs
and woolens, while the cigarette-beetle has
the depraved taste of our modern youth.

v
'
x md
m
FIG. ii. FIG. 12.
FIG. ii. Mouth-parts, much enlarged, of the house-fly, Musca domestica. mx.p., maxil-
lary palpi; lb., labrum; Ii., labium; la., labellum.
FIG. 12. Head and mouth-parts, much enlarged, of thrips. ant., antenna; lb., labrum;
md., mandible; mx., maxilla; mx.p., maxillary palpus; li.p., labial palpus; m.s.,
mouth-stylet. (After Uzel; much enlarged.)

With all this variety of food, it is obvious that the food-taking parts must
show many differences; one insect needs strong biting jaws (Fig. 8), another
a sharp piercing beak (Figs. 9, 13, and 14), another a long flexible sucking

proboscis (Figs. 10 and 16), and another

a broad lapping tongue (Fig. ii).

Just this variety of structure actual y exists, and in it the classific entomolo-
gist has found
a basis for much of his modern classification.
Throughout all this range of mouth structure the insect morphologists
and students of homology, beginning with Savigny in 1816, have be:n able
to trace the fundamental three pairs of oral jointed appendages, the mandi-
bles, maxillae, and labium. Each pair appears in widely differing condi-
tions; the mandibles may be large strong jaws for biting and crushing, as
with the locust, or trowel-like, for moulding wax, as with the honey-bee, or
 The Structure and Special Physiology of Insects 9

long, flat, slender, and saw-toothed, as with the scorpion-flies, or needle-like,

as in all the sucking bugs, or reduced to mere rudiments or wholly lacking,
as in the moths and butterflies. Similarly with the other parts. But by
careful study of the comparative anatomy of the mouth structure, and par-
ticularly by tracing its development in typical species representing the
various types of biting, sucking, and lapping mouths, all the various kinds of
mouth structure can be compared and the homologies or structural cor-
respondences of the component parts determined. Figs. 8 to 16 illustrate

FIG. -13. FIG. 14. FIG. 15.

FIG. 13. Seventeen-year cicada, Cicada septendecim, sucking sap from twig. (After
Quaintance; natural size.)
FIG. 14. Section of twig of Carolina poplar showing beak of cicada in position when
sucking. (After Quaintance; much enlarged.)
FIG. 15. Mouth-parts, much enlarged, of net-winged midge, Bibicocephala doanei,
female, md., mandible; mx., maxilla; mx.L, maxillary lobe; mx.p., maxillary
palpus; li., labium; hyp., hypopharynx; pg., paraglossa of labium; l.ep., labrum
and epi pharynx.

examples of different mouth structures, with the corresponding parts similarly

lettered.
The most conspicuous structural characteristic of insects is their poses-
sion of wings. And the wings undoubtedly account for much of the success
of the insect type. Insects are the dominant animal group of this age, as
far as number of species constitutes dominance, their total largely sur-

passing that of the species of all the other kinds of living animals. Flight
is an extremely effective mode of locomotion, being swift, unimpeded by
obstacles, and hence direct and distance-saving, and an animal in flight
is safe from most of its enemies. The wings of insects are not modified true
appendages of the body, but arise as simple sac-like expansions (Fig. 17)
of the body-wall or skin much flattened and supported by a framework of
io The Structure and Special Physiology of Insects

strongly chitinized lines called veins. These veins are corresponding cutic-
ular thickenings, in the upper and lower walls of
the flattened wing-sac, which protect, while the

wing is forming, certain main tracheal trunks that

carry air to the wing-tissue. After the wing is

expanded and dry, the tracheae mostly die out, and

the veins are left as firm thick-walled branching
tubes which serve admirably as a skeleton or
framework for the thin membranous wings. It

has been found that despite the obvious great

variety in the venation, or number and arrange-
ment of these veins of the wing, a general type-
plan of venation is apparent throughout the insect
class. The more important and constant veins have

been given names, and their branches numbers

(Fig. 1 8). By the use of the same name or
number for the corresponding vein throughout all

the insect orders, the homologies or morphological .

correspondences of the veins as they appear in the

variously modified wings of the different insects
are made apparent Many figures scattered through
this book show the venation of insects of

different orders,and the corresponding

lettering and numbering indicate the
homologies of the veins. As the wing
venation presents differing conditions
readily noted and described, much use is
made of it in classification.
The differences in the wings them-
selves, that is, in number, relative size

of fore and hind wings, and in struc-

ture, i.e., whether membranous and
delicate, or horny and firm, etc., have
FIG. 16.
always been used to distinguish the
FIG. 16. Sphinx moth, showing proboscis; , , .

at left the proboscis is shown coiled up larger groups, as orders, of insects,

on the under side of the head, the nor- ancj tne nrst classification, that of
mal position when not in use. (Large .... , ,. ., . .

Linnaeus (1750 app.), divides the class

figure, one-half natural size; small fi|-
ure, natural size.) into orders almost solely on a basis
of wing characters. The ordinal names expressed, to some degree, the
differences, as Diptera,* two-winged; Lepidoptera, scale-winged; Coleoptera,
sheath- winged, and so on. As a matter of fact, there may be much differ-
* The derivation of the Linnsan ordinal names is given on p. 223.
 The Structure and Special Physiology of Insects 1 1

ence in the wings within a single order; most beetles, for example, have
four wings, but some have two and some none. There are indeed wingless
species in almost every insect order. But a typical beetle has quite dis-
tinctive and commonly recognized wing characters; that is, it has two pairs
of wings, the fore pair being greatly thickened, and developed to serve as
sheaths for the larger, membranous under-pair, which are the true flight
wings. Similarly, practically all moths and butterflies have two pairs of

FIG. 17. FIG. 18.

FIG. 17. Wing of cabbage-butterfly, Pieris rape?, in early sac-like stage, tr., trachea;
//., tracheoles; l.v., lines of future veins. (After Mercer; greatly magnified.)
FIG. 18. Diagram of wings of monarch butterfly, Anosia plexippus, showing venation.
c., costal vein; s.c., subcostal vein; r., radial vein; CM., cubital vein; a., anal veins.
In addition, most insects have a vein lying between the subcostal and radial veins,
called the median vein. (Natural size.)
membranous wings completely covered above and below by small scales,
which give them their distinctive color and pattern.
The exoskeleton, or cuticle, of the insect body is sometimes nearly
smooth and naked, but usually it is sculptured by grooves and ridges, punc-
tures or projections, and clothed with hairs or those modified flattened hairs
known as scales (especially characteristic of butterflies and moths). This
clothing of hairs or scales, or the skin itself, is variously colored and pat-
terned, often with the obvious use of producing protective resemblance or
mimicry, but often without apparent significance. (Fo- an account of the colors
and patterns of insects and their uses see Chapter XVII.) The hairs may serve
for protection, or may be tactile organs, or even organs of hearing (see p. 26).
The projecting processes may be spines or thorns or curious and inexplicable
i 2 The Structure and Special Physiology of Insects

knobs and horns. The rhinoceros-beetle (Dynastes) (Fig. 19) and the sacred
scarabeus are familiar examples of insects with such prominent processes.
The insect body, as a whole, appears in great variety of form and range
of size, as ourknowledge of the variety of habit and habitat of insects would
lead us to expect. In size they vary from the tiny four-winged chalcids
which emerge, after their parasitic immature life, from the eggs of other
insects, and measure less than a millimeter in length, to the giant Phasmids

FIG. 19. Rhinoceros-beetle, Dynastes tityrus, showing chitinous horns.

(walking-sticks) of the tropics, with their ten or twelve inches of body length,
and the great Formosan dragon-flies with an expanse of wing of ten
inches. A Carboniferous insect like a dragon-fly, known from fossils found
at Commentry, France, had a wing expanse of more than two feet.
Insects show a plasticity as to general body shape and appearance that results
in extreme modifications corresponding with the extremely various habits
of life that obtain in the class. Compare the delicate fragility of the gauzy-
winged May-fly with the rigid exoskeleton and horny wings of the water-

beetle; the long- winged, slender-bodied flying-machine we call a dragon-

fly with the shovel-footed, half-blind, burrowing mole-cricket; the plump,

toothsome white ant that defends itself by simple prolificness with the spare,
angular, twig-like body of the walking-stick with its effective protective
resemblance to the dry branches among which it lives. Compare the leg-
less, eyeless, antennaless, wingless, sac-like degraded body of the orange-
scale with the marvelous specialization of structure of that compact expo-
nent of the strenuous insect life, the honey-bee; contrast the dull colors of the
lowly tumble-bug with the flashing radiance of the painted lady-butterfly.
But through all this variety of shape and pattern, complexity and degenera-

tion, one can see the simple fundamental insect body-plan; the successive

segments, their grouping into three body-regions, the presence of segmented

appendages on head and thorax and their absence on abdomen (except
perhaps in the terminal segments), and the modification of these append-
ages into antennae and mouth-parts on the head, legs on the thorax, and
ovipositor, sting, or claspers in the abdomen.
In the character of the structure and functions of the internal organs
 The Structure and Special Physiology of Insects I
3

or systems of organs of insects, a special interest attaches to the conditions

shown by the circulatory and respiratory systems, and by the special sense-

< j2L
i

-^i^- - ;
*
, "**-,,
J _ /;/-?_/.&
"'
i M> '^^LT'^.
*r

00

FIG. 20. Diagram of lateral interior view of monarch butterfly, Anosia plexippus, show-
ing the internal organs in their natural arrangement, after the removal of the right
half of the body-wall together with the tracheae and fat body; I to III, segments
of the thorax; i to 9, segments of the abdomen. Alimentary Canal and Appen-
dages: ph., pharynx; sd. and sgl., salivary duct and gland of the right side; oe.,
oesophagus; }.r., food-reservoir; st., stomach; i., small intestine; c., colon; r., rec-
tum; a., anus; m.v., Malpighian tube. Haemal System: h., heart or dorsal vessel;
ao., aorta; a.c., aortal chamber; Nervous System (dotted in figure): br., brain;
g., suboesophageal ganglion; l.g., compound thoracic ganglia; ag. v ag. v first and
fourth abdominal ganglia. Female Reproductive Organs: cp., copulatory pouch;
v., vagina; o., oviduct, and oo., its external opening; r.ov., base of the right ovarian
tubes turned down to expose the underlying organs; l.ov., left ovarian tubes in posi-
tion, and ov.e., their termination and four cords; sp., spermatheca; a.gl. r part
of the single accessory gland; a.gl. 2 one of the paired accessory glands; only the
,

base of its mate is shown. Head: a., antenna; mx., proboscis; p., labial palpus.
(After Burgess; three times natural size.)

organs and their manner of functioning. The muscular system varies from the
simple worm-like arrangement of segmentally disposed longitudinal and
ring muscles possessed by the caterpillars, grubs, and other worm-like larvae,
to the complicated system of such

specialized and active forms as the

honey-bee and house-fly. Lyonnet
describes about two thousand dis-
tinct muscles in the caterpillar of
the goat-moth. Insect muscles are
similar, in their finer structure, to
those of other animals, most of FIG. 21. Bit of muscle of a biting bird-louse,

them of Eurymetopus taurus. (Greatly magnified.)

being composed finely
cross-striated fibers (Figs. 21 and 22) held together in larger or smaller
masses and attaching to the rugosities of the inner surface of the exo-
skeleton. The muscle substance, when fresh, is peculiarly transparent
and it has great contractile power.
delicate-looking, but
The is a tube
alimentary canal (Figs. 23-27), like that of other animals,
but little longer than the body in flesh-eating forms, and much longer in
it runs, more or less curving and coiled, through the body
plant-feeders;
irom mouth to anal opening, which lies in the last segment of the abdomen.
14 The Structure and Special Physiology of Insects

This tube is expanded variously to form crop, gizzard, or stomach, and

'""

W4-UWWUUUI

Iffiffilll
GIIIlimilfflE'

FIG. 22. Diagrammatic figures of bits of insect muscle, variously treated. (After Van
Gehuchten; greatly magnified.)
contracted elsewhere to be oesophagus or intestine.
One or two pairs of salivary glands pour their fluid into
the mouth, while the digesting stomach or ventriculus

usually possesses two or more pairs of diverticula known

as gastric cceca, which are lined with glands believed
to secrete special digestive fluids. Neither liver
nor kidneys are present in the insect body, but the
secretory function of the latter are undertaken 'by a
number of usually long thread-like tubular diverticula
of the intestine known as Malpighian tubules. The
intestine itself is usually obviously made up of three
successive parts, a large intestine, small intestine,
and rectum. There are also
present not infrequently in-
testinal ca'ca.
Two striking peculiarities - a I.e.
about the reproductive system
of insects are the possession

by the female of one or more

FIG. 23. Alimentary
canal of a locust. At spermathecae (Fig. 66, r.s.) in
upper end the oesoph- which the male fertilizing
agus, then the ex-
cells, the spermatozoa, are re-
panded crop, then sev-
eral large gastric cceca, ceived and held, and the com-
then the true stomach, all the envelopes of
pletion of
the thread-like Malpig- ... . ,
FIG. 24. Dissection of
hian tubules, the bent the egg, including the outer cockroach to show (al.c.)
intestine, and the ex- hard shell, before its specific alimentary canal. (After
panded rectum. (After fertilization
, .,. , !, T-, Hatschek and Cori: twice
takes place. Fer- natural size.)
Snodgrass; enlarged.)
 The Structure and Special Physiology of Insects i
5

tilization is itself accomplished in the lower end of the egg-duct just before the
egg is laid,
by the escape of spermatozoa from the spermatheca (the female

saLg..

..ea.

.mi.

FIG. 25. FIG. 26.

FIG. 25. Alimentary canal of larva of harlequin-fly (Chironomus sp.). oes., oesophagus;
s.g., salivary gland; ca., cardiac chamber of stomach; ml., Malpighian tubules; ch.,
intestinal chamber; 5/., small intestine; col., colon. (After Miall and Hammond;
much enlarged.)
FlG. 26. Alimentary canal of two species of thrips; at left Trichothrips copiosa, male,
at right Aelothrips fasciata. sal.g.,salivary gland; oes., oesophagus; prov., proven-
triculus; vent., ventriculus; m.t., Malpighian tubules; int., intestine; rec., rectum.
(After Uzel; greatly enlarged.)

having of course previously mated) and their entrance into the egg through a
tiny opening, the micropyle (Fig. 67), in the egg-shell and inner envelopes.
A queen bee mates but once, but she may live for four or five years after
this and continue to lay fertilized eggs during all this time. She must
j 6 The Structure and Special Physiology of Insects

receive several million spermatozoa at mating, and retain them alive in the

spermatheca during these after-years.

*
int.

FIG. 27. Alimentary canal of dobson-fly, Corydalis cornuta. A, larva; B, adult; C, pupa;
oes.,oesophagus; prov., proventriculus; g.c., gastric cceca; vent., ventriculus; r.g.,
reproductive gland; m.t., Malpighian tubules; int., intestine; inl.c., intestinal
coecum; rec., rectum; drg., oviduct. (After Leidy; twice natural size.)

The circulatory system of insects presents two particular features of inter-

est in that the blood does not, as in our bodies, carry oxygen to the tissues, and

FlG. 28. Cross-section and longitudinal section- of salivary gland of giant crane-fly,
Holorusia rubiginosa. (Greatly magnified.)

that there is a contractile pulsating heart-like organ, but no arteries or veins.

The so-called heart is a delicate-walled, narrow, subcylindrical vessel com-

posed of a series of most commonly from three to eight successive cham-

bers lying longitudinally along the median line just underneath the dorsal
wall of the abdomen and thorax (Figs. 30 and 31). Each chamber opens,
guarded by a simple valvular arrangement (Fig. 33), into the chambers
 The Structure and Special Physiology of Insects 1 7

behind and before it, the posterior one being closed behind and the anterior

FIG. 29. Cells of digestive epithelium of stomach (ventriculus) of crane-fly, Ptychoptera

sp.,showing secretion of digestive fluids, or expulsion of cell-content. (After Van
Gehuchten; greatly magnified.)

one extending forward into or near the head as a narrowed tubular anterior
portion, which is sometimes called the
aorta. From the anterior open end of
this aorta the blood, forced
by pulsations
of the heart- chambers, which proceed

rhythmically from the posterior one

forward, pours out into the body-cavity,
proceeding in more or less regular cur-
rents or paths, but never enclosed in
arterial vessels, bathing all the tissues,
and carrying food to them. Finally
taking up fresh supplies of food by bath-
ing the food-absorbing walls of the

alimentary canal, it enters the chambers

of the heart through lateral openings in
these (either at the middle or anterior end
of each), which thus establish communi-
cation between the body-cavity and heart- FIG. 30. FIG. 31.
The blood receives no more oxygen 'than FIG. 30.Diagram of circulatory
it needs for its own use, and thus does system of a young dragon-fly; in
middle is the chambered dorsal
not play nearly so complex a function in vessel, or heart, with single artery.
the insect's body as in ours. And this Arrows indicate direction of blood-
currents. (After Kolbe.)
simplicity of function probably explains FIG. 31. Dissection showing dorsal
in some degree the extreme primitiveness vessel, or heart, of locust, Dis-
sosteira Carolina. (After Snodgrass;
of the make-up of the circulatory system.
twice natural size.)
It will be seen that the respiratory
system, on the other hand, is particularly highly developed, as it devolves
1 8 The Structure and Special Physiology of Insects

FIG. 32. FIG. 33-

FIG. 32. Portion of dorsal vessel and pericardial membrane of locust, Dissosteira Caro-
lina. (After Snodgrass; greatly magnified.)
FIG. 33. Cross-section of dorsal vessel or heart in pupa of tussock-moth, Hemerocampa
leucostigma, showing valves. (Greatly magnified.)

at.

FIG. 34. FIG. 35. FIG. 36.

FIG. 34. Diagram of tracheal system in body of beetle,

sp., spiracles; tr., tracheae.
(After Kolbe.)
FIG. 35. Diagram showing main tracheae in respiratory system of locust, Dissosteira
Carolina. (After Snodgrass; twice natural size.)
FIG. 36. Diagram showing respiratory system in thrips. si., spiracles. (After Uzel;
much enlarged.)
 The Structure and Special Physiology of Insects 1
9

on it not merely to take up oxgyen from the outer air and give up the
waste carbon dioxide of the
body, but also to convey these
gases to and from all the tis-

sues of the body. The blood

is not red, but
pale yellowish
or greenish, and is really more
like the lymph of the ver-
tebrate body than like its

blood
Insects do not breathe
through the mouth or any
openings on the head, but have
a varying number (usually
from two to ten pairs) of
small paired openings on the
FIG. 37. FIG. 38. s ides of the thorax and abdo-

FlG. 37. Diagram showing respiratory system of pupa men. These openings, called
of mealy-winged fly, Aleyrodes sp.; only two pairs .
,
s P iracles > O] otiamata
ta are ar '
(After Bemis; much
>
of spiracles are present.
enlarged.) ranged segmentally and in
FIG 38. Diagram head of cockroach. most msects are to be found
of tracheae in
Note branches to all mouth-parts, and the an-
(After Miall on two
tennae, t., trachea, or air-tubes. of the thoracic seg-
and Denny.) ments and on all the abdomi-
nal segments except the last two or three. The openings are guarded by fine
hairs or even little valvular lids to prevent

and are the entrances to

the ingress of dust,
an extended system of delicate air-tubes or
tracheae which branch and subdivide until
the whole of the internal body is reached
and ramified by fine capillary vessels bring-

ing fresh air to all the tissues and carrying

off the waste carbon dioxide made by the
metabolism of these tissues. The usual
general arrangement of this elaborate re-
spiratory system is shown in Figs. 34, 35,
and 36. Short broad trunks lead from
each spiracle to a main longitudinal trunk
on each side of the body, from which
numerous branches arise, these going to
particular regions of the body (Fig. 38) FIG. 39. Piece of trachea (air-tube),
reatl magnified, showing spiral
and there branching repeatedly '
until g, j
. .
thread (taemdia). (Photomicro-
even individual cells get special tiny graph by George O. Mitchell.)
2O The Structure and Special Physiology of Insects

respiratory capillaries.The tracheae are readily recognized under the micro-

scope by their finely transversely ringed or striated appearance (Fig. 39).
These transverse "rings" are really spirally arranged short chitinized
thread-like thickenings on the inner wall of the tube, which by their elasticity

keep the delicate air-tubes open. The tubes are filled and emptied by a
rhythmic alternately contracting and expanding
movement of the called the respiratory
abdomen,
movement. When
the ring-muscles contract, the
walls of the abdomen are squeezed in against
the viscera, which, compressing the soft air-tubes,
force the air out of them through the spiracles;
when the body-walls are allowed to spring back
to normal position fresh air rushes in through the
spiracles and fills up the air-tubes, which expand
because of the elastic spiral thickenings in their
walls. Insects which live in water either come

up to the surface to breathe and in some cases

to take down a supply of air held on the outside
of the body by a fine pubescence like the pile of
velvet, or they are provided with tracheal gills

(Fig. 40) which enable them to breathe the air

mixed with, or dissolved in, the water. Gilled
insects do not, of course, have to come to the
surface to breathe. The gills may be thin plate-
FIG. 40. Young (nymph) of
like flaps on the sides or posterior tip of the
May-fly showing (g.) tra-
cheal gills.(After Jenkins body, or may be tufts of short thread-like tubes
and Or they
Kellogg.) variously arranged over the body.
may be, as in the dragon-fly nymphs, thin folds along the inner wall of the

rectum, the water necessary to bathe them being taken in and ejected again
through the anal opening. In aU cases these insect gills differ from those
of other animals, as crabs and fishes, in that they are not organs for the

purification of the blood, i.e., effecting an exchange of carbon dioxide

and
oxygen carried by it, but are means for an osmotic exchange of the fresh
air dissolved inwater for carbon-dioxide-laden air from air-tubes or tracheae
which run out into the gills. Probably no more blood enters these gills
than is necessary to bring food to them. Impure air is brought to them
by air-tubes, and exchanged by osmosis through the thin walls of air-tube
and gill-membrane for fresh air, which passes from these gill air- tubes to
the rest of the respiratory system of the body.
The nervous system of insects shows the fundamentally segmental make-up
of the body better than any of the other systems of internal organs, although

probably in the successive chambers of the dorsal vessel or heart, and certainly
 The Structure and Special Physiology of Insects 2 1

in the pairedarrangement of the spiracles and tracheal trunks leading from

them, a segmental condition is obvious. The central nervous system consists

FIG. 41. Larva of giantcrane-fly, Holorusia rubiginosa. A, entire; B, dissected, show-

ing organs except the muscles and ventral nerve-chain, h., head; ant., antenna;
all

i.b.res., imaginal bud of pupal respiratory tube; i.b.wg., imaginal bud of wing;
i.b.msj., imaginal bud of mesothoracic leg; i.b.h., imaginal bud of balancer;
i.b.mt.1., imaginal bud of metathoracic leg (the imaginal buds of fore' legs are con-
cealed by head-capsule) sal.gl., salivary gland (the other salivary gland is removed)
; ;

br., brain; ces., oesophagus; prov., proventriculus; susp., suspensorium ; g.c., gastric
coecum; vent., ventriculus; tr., trachea; ad.tis., adipose tissue; mal.lub., Malpi-
ghian tubule; d.v., dorsal vessel; w.m., wing-muscles of pericardium; sm.int.,
small intestine; tes., testis; int.c., intestinal caecum; v.d., vas deferens; Lint., large
intestine; sp., spiracle; term.pr., terminal processes. (Twice natural size.)

of a brainand a ventral chain of pairs of ganglia segmentally arranged and

connected by a pair of longitudinal cords or commissures (Figs. 42, 43, 44).
The two members of each of the pairs of ganglia as well as of the pair of
 22 The Structure and Special Physiology of Insects

FIG. 42. FIG. 43. FIG. 44.

FIG. 42. Diagramof ventral nerve-cord of locust, Dissosteira Carolina. (After Snod-
grass; twice natural size.)
FIG. 43. Diagram of the nervous system of the house-fly.- (After Brandt; much
enlarged.)
FIG, 44. Nervous system of a midge, Chironomus sp. (After Brandt, much enlarged.)

commissures are in most insects more or less fused to form single ganglia
and a single commissure, but in others the commissures,
at least, are quite distinct. In the simpler or more
generalized condition of the nervous system as seen
in the simpler insects and the larvae of the higher
ones there are from three or four to seven or eight
abdominal ganglion pairs, one pair to a segment, a
pair in each of the three thoracic segments, and one
in the head just under the oesophagus. From this
ganglion (or fused pair) circumcesophageal commis-
sures run up around the oesophagus to an important
FIG. 45. Brain, com- ganglion (also composed of the fused members of a
pound and part
eyes, pair) lying just above the oesophagus and called the
of sympathetic nerv-
ous system of locust, brain, or supracesophageal ganglion (Figs. 45, 46, and
Dissosteira Carolina.
47). From this proceed the nerves to those impor-
(After Snodgrass;
on the head, the
tant organs of special sense situated
greatly magnified.)
antennae and eyes. From the subcesophageal gan-
glion nerves run to the mouth-parts, from the thoracic ganglia to the
 The Structure and Special Physiology of Insects
23

wings and legs and the complex thoracic muscular system, while from
the abdominal ganglia are innervated the abdominal muscles and sting,
ovipositor, or male claspers. In addition to this main or ventral nervous
system there is a small and considerably varying sympathetic system (Figs.
46 and 48) to which belong a few minute ganglia sending nerves to those
viscera which act automatically or by reflexes, as the alimentary canal and
heart. This sympathetic system is connected with the central or principal

FIG. 46. FIG. 47.

FIG. 46. Brain, circumoesophageal commissures, and subcesophageal ganglion of the

red-legged locust, Melanoplus jemur-rubrum. oc., ocellus; op.n., optic nerve; a.n.,
antennal nerve; m.oc., middle ocellus; op. I., optic lobe; a.l., olfactory lobe; a.s.g.,
anterior sympathetic ganglion; p.s.g., posterior sympathetic ganglion; f.g., frontal
sympathetic ganglion; Ibr., nerve to labrum; oe.c., circumcesophageal commissure;
g*, suboesophageal ganglion; md., nerve to mandible; mx., nerve to maxilla; /..,
nerve to labium; n., unknown nerve, perhaps salivary. (After Burgess; greatly
magnified.)
FIG. 47. Cross-section of brain, cesophagus, circumcesophageal commissures, and
suboesophageal ganglion of larva of the giant crane-fly, Holorusia rubiginosa.

nervous system by commissures which meet the brain just at the origin
from it of the circumcesophageal commissures.
The specialization of the ventral nerve-chain is always of the nature of
a concentration, and especially cephalization of its ganglia (Figs. 49 and
50). The abdominal ganglia may be fused into two or three or even into
one compound ganglion; or indeed all of them may migrate forward and
fuse with the hindmost thoracic ganglion, thus leaving the whole abdomen
24 The Structure and Special Physiology of Insects

to be innervated by long nerves running from the thorax. The thoracic

ganglia may fuse to form one, and in extreme cases all the abdominal and
thoracic ganglia may be fused into one large mid-
thoracic center.
In tracing the development of the nervous
system during the ontogeny of one of the special-
ized insects, the changes from generalized condi-
tion, i.e., presence of numerous distinct ganglia

segmentally disposed, shown in the newly hatched

FIG. 48. FIG. 49.

FlG. 48. Part of sympathetic nervous system of larva of harlequin-fly, Chironomus

dorsalis. oes., oesophagus; j.g., frontal ganglion; r.n., recurrent nerve; d.v., dorsal
4
vessel; n , nerve passing from brain to frontal ganglion (Newport's fourth nerve);
brain; rn., point of division of recurrent nerve; tr., tracheae; pg., paired ganglia;
br.,
nerve of dorsal vessel; d.v.g., ganglia of dorsal vessel; g.n., gastric nerve
d.v.n.,
to cardiac chamber. The course of the recurrent nerve beneath the dorsal vessel is
dotted. (After Miall and Hammond; greatly magnified.)
FIG. 49; Stages in the development of the nervous system of the honey-bee, Apismelli-
fica; i showing the ventral nerve-cord in the youngest larval stage, and 7 the system
in the adult. (After Brandt; much enlarged.)

i.e., extreme concentration and cephalization,

larva, to specialized condition,
that migration
is, forward and fusion of the ganglia, shown in the adult,
are readily followed (Figs. 49 and 50).
The special senses of insects and the sense-organs are of particular inter-
est because of the marked unusualness of the character of the specialization
of both the organs and senses, as compared with the more familiar condi-
tions of the corresponding organs and functions of our body. The world
is known to animals only by the impressions made by it on the sense-organs,
 The Structure and Special Physiology of Insects 25
and the particular condition of functioning of these organs, therefore, is of

unique importance any particular animal. If the senses vary

in the life of
much in their capacities among different animals, the world will have a differ-
ent seeming to different creatures. It will be chiefly known to any par-

ticular species through the dominant sense of that species. To the con-
genitally blind the world is an experience of touched things, of heard things,
and of smelled and tasted things. To the bloodhound it is known chiefly
by the scent of things. It is a world of odors; the scent of anything deter-
mines its
dangerousness, desirableness, its interestingness.
its As insects
know it, then, the world depends largely upon the particular character and
capacity of (heir sense-organs, and we realize on even the most superficial
examination of the structure of these organs, and casual observation of the

FIG. 50. Stages in the development of the nervous system of the water-beetle, Mcilius
sulcatus; i showing the ventral nerve-cord in the earliest larval stage, and 7 the
system in the adult. (After Brandt; much enlarged.)

responses of insects to those stimuli, like sound-waves, light-waves, dis-

solved and vaporized substances, which affect the sense-organs, that the
insects have some remarkable special sense-conditions. But the difficul-
ties in the
way of understanding the psychology of any of the lower animals
are obvious when it is recalled that our only knowledge of the character
of sense-perceptions has to depend solely on our experience of our own per-

ceptions, and on the basis of comparison with this. We

do not know if
hearing is same phenomenon or experience with insects as with us.
the
But a comparison of the morphology of the insect sense-organs with that
of ours, and a course of experimentation with the sight, hearing, smelling,

etc., of insects, based on similar experimentation with our own senses, leads
us to what we believe is some real knowledge of the special sense-condi-
tions of insects.
2.6 The Structure and Special Physiology of Insects

Insects certainly have the senses of touch, hearing, taste, smell, and sight.
If they have others, we do not know it, and probably cannot, as we have
no criteria for recognizing others.
The tactile sense resides especially
in so-called
"
tactile hairs," scattered
more or less abundantly or regu-
larly over thebody. Each of these
hairs has at its base a ganglionic
nerve-cell from which a fine nerve
runs to some body ganglion (Fig. 51).
They are specially numerous and

conspicuous on the antennae or

of a "
FIG. 51. Diagram showing innervation feelers," and often on certain pro-
tactile hair, sh., tactile hair; ch., chitinized ,, , . . ,
.,

cuticle;hyp., hypoderm, or cellular layer

cesses called cerci > projecting from
of the skin; s.c., ganglion cell; c.o., gan- the tip of the abdomen. They may
CCntral nerV US SyStCm (AftCr
OCCUr however on an y P art of the
'

' >
vo^Raih)
body, and are usually recognizable
by length and semi-spinous nature.
their The sense of taste resides
in certain small papillae, usually two-segmented, or in certain pits, which

FIG. 52. FIG. 53.

FIG. 52. Nerve-endings in maxillary palpus of Locusta viridissima. s.h., sense-
tip of
hairs; s.c., sense-cells; b.c., blood-cells. (After vom Rath; greatly magnified.)
FIG. 53. Nerve-endings in tip of labial palpus of Machilis polypoda. (After vom
Rath; greatly magnified.)
occur on the upper wall of the mouth (epipharynx) and on the mouth-
parts, especially the tips of the maxillary and labial palpi, or mouth-
feelers. As substances to be tasted have to be dissolved, and have to
 The Structure and Special Physiology of Insects 27

come into actual contact with the special taste nerves, it is obvious
that insects, to taste solid foods, have first to dissolve particles of these

foods in the mouth-fluids, and that the taste-organs have to be situated

in the mouth or so that they can be brought into it to explore the food, as
are the movable, feeler-like palpi. What experimentation on the sense of
taste in insects has been carried on shows that certain insects certainly taste
food substances, and indicates that the sense is a common attribute of all
insects. Lubbock's many experiments with ants, bees, and wasps present
convincing proof of the exercise of the taste sense by these insects. Forel
mixed morphine and strychnine with honey, which ants, attracted by the
honey smell, tasted and refused. Will's experiments show that wasps

recognize alum and quinine by taste. He found bees and wasps to have
a more delicate gustatory sense than flies.
Smell is probably the dominant special sense among insects. It exists
at least in a degree of refinement among certain forms that is hardly

equalled elsewhere in the animal kingdom. The smelling organs are micro-
scopic pits and minute papillae seated usually and especially abundantly
on the antennae, but probably also occurring to
some extent on certain of the mouth-parts. The
fact that the antennae are the principal, and in

many insects the exclusive, seat of the olfactory

organs has been proved by many experiments in

removing the antennae or coating them with par-
affine. Insects thus treated do not find food or
each other. As substances to be smelled must
actually come into contact, in finely divided con-

dition, with the olfactory nerve-element, these

pits and papillae are arranged so as to expose
the nerve-end and yet protect it from the
ruder contact with obstacles against which the
antenna; may strike. It is certain that most
insects find their food by the sense of smell, and
the antenna of a carrion-beetle (Fig. 54) shows

plainly the special adaptation to make this sense

highly effective. On the "leaves" of each antenna
FIG. 54. Antenna of a
of June-beetles nearly 40,000 olfactory pits occur.
carrion-beetle with- the
Some of the results of experimentation on smell terminal three segments
indicate a delicacy and enlarged and flattened,
specialization of this sense and bearing many smell -
hardly conceivable. A few examples will illustrate ing-pits. (Photomicro-
graph by George O. Mit-
this. way back
It is believed that ants find their
chell; much enlarged.)
to their by the sense of smell, and that
nests

they can recognize by scent among hundreds of individuals taken from

 28 The Structure and Special Physiology of Insects

various communities the members of their own community. Miss Fielde's

experiments show that the recognition of ants by each other depends on the
existence of a sense of smell of remarkable differentiative capacity. The
odors of the nest, of the species, of the female parent, and of the individ-
ual are all distinct and perceivable by the smelling-organs, situated on
distinct particular antennal segments. In the insectary at Cornell University
a few years ago a few females of the beautiful large promethea moth were
put into a covered box which was kept inside of the insectary building.
No males of this moth species had been seen about the insectary nor in
its immediate vicin-

ity for several days,

although they had

been specially sought
for by collectors.
Yet in a few hours
after the female
moths were first con-
fined nearly fifty
male prometheas
were fluttering about
outside over the glass
roof of the insectary.

They could not see

the females, but un-

doubtedly discovered
them by the sense of
smell. These pro-
methea moths have
FIG. 55. Auditory organ of a locust, Melanoplus sp. The elaborately branched
large clear part in the center of the figure is the thin tym-
with the or feathered anten-
panum auditory vesicle (small, black, pear-shaped
spot) and auditory ganglion (at left of vesicle and connected nae, affording area
with it by a nerve) on its inner surface. (Photomicrograph
for very many smell-
by George O. Mitchell; greatly magnified.)
ing-pits.
Mayer's experiments with promethea also reveal the high specialization
of the sense of 'smell. This investigator carried 450 promethea cocoons
from Massachusetts to the Florida keys. Here on separated small
islands the moths issued from the cocoons, hundreds of miles south of their
natural habitat. This isolation insured that no other individuals than
those controlled by the experimenter could confuse the observations.
Female moths were confined in glass jars with the mouth closed by
netting. Other females were confined in smaller glass jars turned upside
down and the mouth buried in sand. Males being released at various
 The Structure and Special Physiology of Insects 29

distances soon found their way to the jar (containing females) which had
its mouth open to the air, but no male came to the jar with its mouth her-

metically sealed. Through the glass sides of both

jars the females were plainly visible. The antennae
of certain males were covered with shellac. These
males, when released, never found the females, and
often paid no attention to them when brought within
an inch of their bodies. Of other males the eyes
were covered with pitch; but these males had no
difficulty whatever in finding the females. It is

plainly obvious from these experiments that the

males found the females wholly by scent and not at
all by sight.
That some insects hear is proved by their posses-
sion of auditory organs, and has also been demon- FIG. 56. Male mosquito,
a
The
strated by experiment. fact, too, that many J^8 gJg r $3S!
insects have special sound-making apparatus and and Kellogg; three times
natural size.)
do make characteristic sounds is a kind of
proof
that they can also hear. The auditory organs of insects, curiously enough,
are of several kinds and are situated on different parts of the body, in
various species. Among the locusts,

katydids, and crickets, the most con-

spicuous of the sound-making in-
all

sects except the cicada, the ears are

small tympanic membranes on the
base of the abdomen in the locusts'
(Fig. 55), and on the tibiae of the fore

legs in the
katydids and crickets.

Associated with each tympanum is a

small liquid-filled vesicle and a special
auditory ganglion from which an
auditory nerve runs to one of the

ganglia of the thorax. Among the

FIG. 57 .-Dia g ram of longitudinal section midges and mosquitoes the antennas-
through first and second antennal seg- those all-important sensitive structures
ments of a mosquito, Mochlonyx culici-
formis, male, showing complex auditory
-are
, , ,, -A A 4 u

abundantly provided With cer-

-

organ composed of fine chitinous rods, tain fine long hairs, the auditory hairs
nerve-fibers and nerve-cells.
Child; greatly magnified.)
(After
(F ;
> j fi)/> which l&ke r ^
soun(j.
waves and transmit the vibrations to an
elaborate percipient structure composed of many fine chitin-rods and ganglion-
ated nerves contained in the next to basal antennal segment (Fig. 57). From
this segment runs a principal auditory nerve to the brain. Many other insects
30 The Structure and Special Physiology of Insects

besides the midges and mosquitoes possess this type of auditory organ;
in fact such an organ, more or less well developed, has been found in almost
every order except the Orthoptera (the order of locusts, crickets, katydids,
etc.) in which the tympanic auditory organs occur.

Special isolated hairs scattered sparsely over the

body, connected with a special peripheral nervous
arrangement, are believed by some entomologists
lo be a third kind of auditory structure, and are
called chordotonal organs. Experimentally the
sense of hearing has been surely determined for
certain insects. A
single striking example of this
experimentation must here suffice. Mayer fastened
a live male mosquito to a glass slide, put it under
FIG. 58. Longitudinal sec-
a microscope, and had a series of tuning-forks of
tion through ocellus of the different pitch sounded. When the Ut 4 fork of
honey-bee, Apis mellifica.
/.,cuticular lens; i.e., cell- 512 vibrations per second was sounded many of
ular layer of skin; c.b., the antennal hairs were set, sympathetically, into
crystalline layer; r.c., ret-
inal cells; o.n.,
strong vibration. Tuning-forks of pitch an octave
optic
nerve. (After Redikor- lower and an octave higher also caused more
zew; greatly magnified.)vibration than any intermediate notes. The male
mosquito's auditory hairs, then, are specially fitted to respond to, i.e., be
stimulated by, notes of a pitch produced by 512 vibrations. Other, but
fewer, hairs of different length vibrated in response to other tones. Those
auditory hairs are most affected which are at right angles to the direction
from which the sound comes. From this it is obvious that, from the position
of the antennae and the hairs, a sound will be loudest or most intense if it is

directly in front of the head. If the mosquito is attracted by sound, it will

thus be brought straight head end on toward the source of the sound. As a

FIG. 59. Ocellar lens of larva of a saw-fly, Cimbex sp., showing its continuity with the
chitinized cuticle. (After Redikorzew; greatly magnified.)

matter of fact, Mayer found the female mosquito's song to correspond nearly
to Ut 4 and
,
that her song set the male's auditory hairs into vibration. With
little doubt, the male mosquitoes find the females by their sense of hearing.
have two kinds of eyes, simple and compound.
Insects On most
species both kinds are found, on some either kind alone, and in a few no
eyes at all. Blind insects have lost the eyes by degeneration. The most
 The Structure and Special Physiology of Insects 3 1

primitive living insects, Campodea and others, have eyes, although only
simple ones. The larvae of the specialized
insects, i.e., those with complete metamor-
phosis, also have only simple eyes. The com-
pound eyes are not complex or specialized
derivations of the simple ones, but are of in-

dependent origin and of obviously distinct

structural character. The simple eyes, also
called ocelli (Fig. 58), which usually occur to
the number of three in a little triangle on

top of the head, are small and inconspicuous,

FIG. 60. Part of corneal cuti- and consist each of a lens, this being simply
cle, showing facets, of the a srna ll convexly thickened clear part of the
compound eye of a horse- , ^,. ., n /T , -,.

fly, Therioplectes sp. (Photo-

chitimzed cuticle of the head-wall (Fig. 59)
micrograph by George O. and a group of modified skin-cells behind it
ell; greatly magnified.)
spedally provi de d with absorbent pigment and
capable of acting as a simple light-sensitive or retinal
surface. The ocellus is supplied with a special nerve
from the brain. The compound eyes are always
paired and situated usually on the dorso-lateral parts
of the head; they are usually large and conspicu-

ous, sometimes, as in the dragon-flies and horse-

flies, even forming two-thirds or more of the mass
of the head. Externally each compound eye pre-
sents a number (which varies all the way from a
score to thirty thousand)
microscopic of facets or

polygonal cuticular windows (Fig. 60). These are

the cornea of the eye. Behind each facet is a dis-
tinct and independent subcylindrical eye-element or
ommatidium composed of a crystalline cone (want-

ing in many insects) enveloping pigment (which pre-

sumably excludes all light-rays except those which
fall perpendicularly or nearly so to the corneal
lens ofparticular ommatidium), and a slender
that

tapering part including or composed of the nervous FIG. 6 r. Longitudinal

or retinal element called rhabdom (Fig. 61). Each section through a few
of these ommatidia perceives that bit of the external
(ornmatidia)of the
object which is directly in front of it; i.e., from which compound eye of a
th '

light is reflected perpendicularly to its corneal facet. .

/^ ^Ttaufne
All of these microscopic images, each of a small part cones; p., pigment; r.,
retinal parts ;<>. optic
of the external object,
J
form a mosaic of the whole
nerve. (After Exner;
object, and thus give the familiar name mosaic greatly magnified.)
32 The Structure and Special Physiology of Insects

vision to the particular kind of seeing accomplished by the compound

eye.
The character or degree of excellence of sight by the two kinds of

eyes obviously varies much. The fixed focus of the ocelli is extremely short,

OiOQ

FIG. 62.

II

FIG. 64. FIG. 63.

FlG. 62. Longitudinal sections through outer part of eye-elements (ommatidia) of com-
pound eyes of Lasiocampia quercijolia; ommatidia at left showing disposition of
pigment in eyes in the light, at right, in the dark. (After Exner; greatly magnified.)
FlG. 63. Longitudinal section through a few eye-elements of the compound eye of Cato-
cola nupta; left ommatidia taken from an insect killed in the dark, right ommatidium
taken from insect killed in the light. (After Exner; greatly magnified.)
FlG. 64. Section through the compound eyes of a male May-fly, showing division of
each compound eye into two parts, an upper part containing large eye -elements
(ommatidia), and a lower part containing small eye-elements (ommatidia). (After
Zimmerman; greatly magnified.)

and probably the range of vision of these eyes is restricted to an inch or

two in front of the insect's head. Indeed entomologists commonly believe
that the ocelli avail little beyond distinguishing between light and darkness.
With the compound eyes the focus is also fixed, but is longer and the range
of vision must extend to two or three yards. It is obvious that the larger
 The Structure and Special Physiology of Insects 33

and more convex the eyes the wider will be the extent of the visual field,
while the smaller and more abundant the facets the sharper and more dis-
tinct will be the image. Although no change in focus can be effected, cer-
tain accommodation or flexibility of the seeing function is obtained by the
movements of the pigment (Figs. 62 and 63) tending to regulate the amount
of light admitted into the eye (as shown by Exner), and by a difference in size
and pigmental character of the ommatidia (Fig. 64) composing the com-
pound eyes of certain insects tending to make part of the eye especially

FIG. 65. A section through the compound eye, in late pupal stage, of a blow-fly, Calli-
phora sarracenice. In the center is the brain with optic lobe, and on the right-hand
margin are the many eye-elements (ommatidia) in longitudinal section. (Photomi-
crograph by George O. Mitchell; greatly magnified.)

adapted for seeing objects in motion or in poor light, and another part for

seeing in bright light and formaking a sharper image (as shown by Zim-
merman for male May-flies, and by myself for certain true flies (see p. 318)).
Our careful studies of the structure of the insect eye, and the experimentation
which we have been able to carry on, indicate that, at best, the sight of
insects cannot be exact or of much range.
The psychology of insects, that their activities and behavior as deter-
is,

mined by their reflexes, instincts, and intelligence, is a subject of great inter-

est and attractiveness, but obviously one difficult to study exactly. The
34 The Structure and Special Physiology of Insects

elaborateness of many insect instincts, such as those of the ants, wasps, and
bees, to choose examples at once familiar and extreme in their complexity,
makes it very difficult to analyze the trains of reactions into individual ones,
and to determine, if it is indeed at all determinable, the particular stimuli
which act as the springs for these various reactions. The attitude of the
modern biologist in this matter would be to keep first in mind the theory
of reflexes, to look keenly for physico-chemical explanations of the reac-

tions, and only when forced from this position by the impossibility of find-

ing mechanical explanations for the phenomena to recognize those com-

plex reflexes which we call instincts, and finally those acts which we call
intelligent, or reasonable, and which are possible only to the possessors of
associative memory. The investigations, mostly recent, which have been
directed toward a determination of the immediate springs or stimuli of
insect reactions indicate clearly that many of these responses, even some
which were formerly looked on as surely indicative of considerable intelli-
gence on the part of their performers, are explicable as rigid reflex (mechan-
ical) reactions to light, gravity, the proximity of substances of certain
chemical composition, contact with solid bodies, etc. On the other hand
the position of the extreme upholders (Bethe, Uexkull, and others) of the

purely reflex explanation of all insect behavior will certainly prove untenable.
As one of the phases of insect biology to which this book is particularly
devoted is that which includes the study of habits, activities, or behavior,

we may dispense with any special discussion of instinct in this introductory

chapter. It is sufficient to say that no other class of invertebrate animals

presents such an interesting and instructive psychology as the insects.

 CHAPTER II
DEVELOPMENT AND META-
MORPHOSIS
HAT animals are born or hatch from eggs in
an immature condition is such familiar natural
history that we are likely to overlook the
significance and consequences of the fact unless
our attention is particularly called to them.
This condition of immaturity makes it necessary
that part of the free life of the organism has
to be devoted to growth and development and
has to be undergone in an imperfect condition,
a condition of structure and physiology, indeed, which may be very different
from that of the parents or of maturity. While most animals that are born
alive resemble the parents in most respects, always excepting that of size,

many of those animals which hatch from eggs deposited outside the body
of the mother issue from the egg with few indeed of the characteristics of the
parents and may be so dissimilar from them that only our knowledge of
the life-history of the animal enables us to recognize these young individuals
as of the samespecies as the parent. The butterfly hatching as the worm-
and the frog as the fish-like tadpole, are the classic examples
like caterpillar,
of this phenomenon. The mammals, our most familiar examples of animals
which give birth to their young alive and free, nourish, for weeks or months
before birth, the developing growing young. But with egg-laying animals
usually only such nourishment is furnished the young as can be enclosed
as food-yolk within the egg-shell. As a matter of fact, some young which
hatch from eggs, as, for example, chickens, quail, etc., hatch in well-
developed condition; and some young mammals, nourished by. the mother's
body until birth, are in a conspicuously undeveloped state, as a young

kangaroo or opossum. But nevertheless it is generally true that an animal

hatched from an egg has still a larger amount of development to undergo
before it comes to the stature and capacity of its parents than one which is
35
 Development and Metamorphosis
born having passed a considerable time growing and developing
alive, after
in the body And this difference in degree of development at
of the mother.
birth is largely due simply to the difference in amount of nourishment
which can be afforded the young. The embryo in the egg uses up its food
early in its developmental career and before it has reached the stage of
likeness to its parents. It issues in a condition picturing some far-distant
ancestor of its species, or morefrequently, perhaps, in a modified, adapted
condition, fit to make of this tender unready creature thus thrust before
itstime into the struggle for living an organism capable of caring for itself,
although not yet endowed with capacities as effective as, or even similar to,
those of the parent.
It is familiar to us, then, that development is not wholly postnatal or

postembryonic that before birth or hatching a greater or less amount of

;

development, requiring a longer or shorter

period of time, has already been undergone.
Every animal begins life as a simple cell; all
animals except the Protozoa (the simplest ani-
mals, those whose whole body for its whole
life is but a single cell) finish life, if red
Nature permits them to come through myriad

dangers safely to maturity, as a complex of

thousands or millions of cells united into
great of tissues and
variety organs. This
od.
great change from most simple to most complex
condition constitutes development: the actual
increase of body-matter and extension of
dimensions is growth.
Most insects hatch from eggs; being born
FIG. 66. Ovaries and oviducts
of a thrips. o.t., ovarial tubes; alive is the exceptional experience of the young
o.d., oviduct; r.s., seminal of but few kinds, and even this is a sort of
receptacle, or spermatheca;
d.r.s., duct of the seminal re- pseudo-birth. Such hatch alive, one may better
ceptacle. (After Uzel; much say, for they begin life in eggs, not laid out-
enlarged.) side the mother body to be sure, but held in
the egg-duct until hatching-time. With very few exceptions, young insects
are not nourished by the mother except in so far as she stores a supply of

yolk around or by the side of each embryo inside the egg-shell. The form-
is a matter which does not lend itself
ing of the egg readily to the observa-
tionand study of amateurs, but is a phenomenon of unusual interest to
whomever is privileged to discover it. The insect ovaries consist of a pair
of little compact groups of short
tapering tubes (Fig. 66). In the anterior or
beginning end of each tube is a microscopic space or chamber from whose
walls cells loosen themselves and escape into the cavity. These cells become
 Development and Metamorphosis 37
either the germinal or the food part of the eggs. There seems to exist no
differentiation among these cells at first, but soon certain ones begin to
move slowly down through the egg-tube in single file, each becoming sur-
rounded and enclosed by yolk, i.e., reserve foodstuff. This gathering of
yolk increases the size of the forming eggs, so that they appear as a short
string of beads of varying size enclosed in the elastic egg-tube. When of
considerable size each egg in the lower end of the tube becomes enclosed

FlG. 67. Insect eggs and parts of eggs, showing micropyle. a, egg of Drosophila cel-
laris; b, upper pole of egg of robber-fly, Asilus crabriformis ; c, upper pole of egg
of hawk -moth, Sphinx populi; d, egg of head-louse, Pediculus capitis; e, egg of
dragon-fly, Libellula depressa; /, upper surface of egg of harpy-moth, Harpyia
vinula; g, upper pole of egg of Hammalicherus cerdo; h, upper pole of egg of sul-
phur-butterfly, Colias hyale. (After Leuckart; much enlarged.)

in two envelopes, a membranous inner one (yolk or vitelline membrane) and

an outer horny one, the chorion or egg-shell. But both of these envelopes
are pierced at one pole by a tiny opening, the micropyle (Fig. 67), and

through this opening the fertilizing spermatozoa enter the egg from the
seminal receptacle just before the egg is extruded from the body.
The development of the embryo within the egg is also securely sealed

away from the eyes of most amateurs. The study of insect embryology
requires a knowledge of microscopic technic, and facilities for fixing and
38 Development and Metamorphosis

imbedding and section-cutting which are not often found outside the college
laboratory. But the particularly interesting and suggestive stages in this
development may be outlined and illustrated in brief space. First, the
germinal cell near the center of the egg divides repeatedly (Fig. 68 A ) and
the resulting new cells migrate outward against the inner envelope of the

egg and arrange themselves here in a single peripheral layer, called the
blastoderm (Fig. 68 D, bT). On what is going to be the ventral side of the
egg the cells of the blastoderm begin to divide and mass themselves to form
the ventral plate (Fig. 69 C). The embryo is forming here; the rest of the
blastoderm becomes modified and folded to serve as a double membranous
envelope (called amnion and serosa) for the embryo. Stretching nearly from
pole to pole as a narrow streak along the ventral aspect of the egg, the

FlG. 68. Early stage in development of egg of water-scavenger beetle, Hydrophilus sp.
A, division of nucleus; B, migration of cleavage-cells outward; C, beginning
first
of blastoderm; D, blastoderm; y., yolk; dc., cleavage-cells; yc., yolk-cells; bl.,
blastoderm. (After Heider; greatly magnified.)

developing embryo begins soon to show that fundamental structural charac-

teristic of insects, a segmental condition (Fig. 6gD). One can now make
out the forming body-rings or segments, and each soon shows the beginnings
or rudiments of a pair of appendages (Fig. 69 E). The appendages of the
head and thoracic segments continue to develop and begin soon to assume
and legs, but those of the
their definitive character of antenna?, mouth-parts,
abdominal segments never get farther than a first appearance and indeed
soon disappear. In the mean time the internal systems of organs are grad-
ually developing, the ventral nerve-chain first, then the alimentary canal,
and later the muscles, tracheae, and the heart. All the time the yolk is
being gradually used up, fed on, by the cells of the developing and growing
embryo, until finally comes the disappearance of all the stored food, and the
time for hatching.
 Development and Metamorphosis 39

The eggs have been laid, because of the remarkable instinct of the
mother, in a situation determined chiefly by the interests of the young
which are to hatch from them. The young of many kinds of insects take
very different food from that of the mother a caterpillar feeds on green
leaves, the butterfly on flower-nectar or live under very different circum-
stances young dragon-flies and May-flies live under water, the adults in
the A monarch butterfly, which does not feed on leaves, nor has ever
air.

before produced young, seeks out a milkweed to lay its eggs upon. The
young monarchs, tiny black-and-white-banded caterpillars, feed on the

FIG. 69. Early stages in the development of the egg of saw-fly, Hylotoma beriberidis.
C, ventral plate removed from egg; D, ventral plate, showing segmentation of body;
E, embryo, showing developing appendages; F, same stage, lateral aspect; G, older
3
stage, lateral aspect, antenna; md., mandible; mx., maxilla; li., labium; /', P, /
ant., ,

legs; sg., St., spiracles; ab.ap., abdominal appendages; n.c., nerve-

salivary glands;
centers; a., anal opening; lb., labrum; sd., cesophageal invagination; y., yolk;
b.s., abdominal segments; pd., intestinal invagination; am., amnion; s., serosa.
(After Graber; greatly magnified.)

green milkweed leaf-tissue; indeed they starve to death if

they cannot have
leaves of precisely this kind of plant! The reason that the butterfly, whose
only food is the nectar of almost any kind of flower, ranges wide to find a
milkweed for its eggs, is one not founded on experience or teaching or rea-
son, but on an inherited instinct, which is as truly and as importantly an
attribute of this particular species of butterfly as its characteristic color

pattern or body structure. And the female of the great flashing strong-
winged dragon-fly, queen insect of the air, when egg-laying time comes,
feels a strange irresistible demand to get these eggs into water, dropping
them in from its airy height, or swooping down to touch the tip of the abdo-
4o Development and Metamorphosis
men to the water's surface, there releasing them, or even crawling down
some water-plant beneath the surface and with arduous labor thrusting the
eggs into the heart of this submerged plant-stem. From the eggs hatch
wingless dwarf-dragons of the pond bottom, with terrible extensile, clutch-

ing mouth-parts and an insatiable hunger for living prey.

So our young insects, after
completing their embryonic development,
come to the time of their
appearance as free individuals compelled to find
their own food and no longer sheltered by a firm egg-shell from the strenu-

eer

FIG. 70. Series of stages in development of egg of fish-moth, Lepisma sp. A, begin-
ning embryo; B, embryo showing segmentation; C, embryo showing appendages;
D, embryo more advanced; E, embryo still more advanced; F, embryo still older
and removed from egg; G, embryo removed from egg at time of readiness to hatch.
y., yolk; emb., embryo; ser., serosa; am., amnion; ant., antenna; lb., labrum;
md., mandible; mx., maxilla; mx.p., maxillary palpus; //., labium; li.p., labial
2 3
palpus; /'.. I
, I, legs; pr., proctodaeum, or intestinal invagination; cer., cercij mp.,
middle posterior process. (After Heymons; greatly magnified.)

ous fighting and hiding of the open road. Now these young insects, depend-
ing upon how far they have carried their developmental course in the egg,
hatch either almost wholly like their parents (excepting always in size), or
in a condition fairly resembling the parents, but lacking all traces of wings
and showing other less conspicuous dissimilarities, or finally they may appear
in guise wholly unlike that of their parents, in such a condition indeed that

they would not be recognized as insects of the same kind as the parents.
But in all cases the young are certain, if they live their allotted days or weeks
 >

Development and Metamorphosis 41

or months, to attain finally the parent structure and appearance. This

attainment is a matter of further development, of postembryonic develop-
ment, and the amount or degree of this development or change is obviously
determined by the remoteness or nearness of the young at the time of hatch-
ing to the adult or parental condition. The young of many of our most
familiar insects, as beetles, flies, moths and butterflies, and ants, bees, and
wasps, hatch out extremely unlike their parents in appearance: the well-
known worm-like caterpillars of butterflies and moths are striking examples
of this unlikeness. The changes necessarily undergone in the develop-
ment from caterpillar to butterfly are so great that there actually results
a very considerable degree of making over, or metamorphosis of the insect,
and for convenience of roughly classifying insects according to their develop-
ment, entomologists have adopted the terms complete metamorphosis,
incomplete metamorphosis, and no metamorphosis to indicate three not
very sharply distinguished kinds or degrees of postembryonic development.
In the latter category are comparatively few species, because most insects
have wings, and no insect is winged But the members
at birth. of the sim-

plest order (Aptera) are all primitively wingless, and their

young are, in practically all particulars except body size and
the maturity of the reproductive glands, like the adults
(Fig. 71) ;development may fairly be said to take place
their
without metamorphosis. In addition to these primitively
simple insects there are certain degenerate wingless species
for example, whose young also
like the biting bird-lice,
reach the parental stature and character without meta-
morphosis.
In the next category, that of development with in-
complete metamorphosis, are included two large orders
of insects and several smaller ones. All the sucking-bugs FIG. 71. Young
(order Hemiptera) and
the locusts, katydids, crickets,
all and adult of p -

and cockroaches (composing the order Orthoptera), as well tne simplesTin-

as the May-flies, dragon-flies, white ants, and several other sects, showing
small groups of unfamiliar forms, agree in having their ?
wf hout 'meta-
t

young hatched in a condition strongly resembling the morphosis.

(Much enlarged.)
parents, although lacking wings, and in some cases, particu-
larly those in which the young live on different food and in a different habitat
from the adults, differing rather markedly in several superficial characters.
Such is the case, for example, with the dragon-flies, whose young are aquatic
and breathe by means of tracheal gills, and are provided with specially con-
structed seizing and biting mouth-parts. But in such essential character-
istics as number of legs, character of eyes and antennae, and, usually, char-

acter of mouth-parts, the young and parent agree. During postembryonic

 Development and Metamorphosis

FlG. 72. Developing stages, after hatching, of a locust, Melanoplus femur-rubrunti

a, just hatched, without wing-pads; b, after first moulting; c, after second moulting,
showing beginning wing-pads; d, after third moulting; e, after fourth moulting,
/,adult with fully developed wings. (After Emerton; younger stages enlarged;
adult stage, natural size.)

FlG. 73. Stages in development of the wings of a locust. /., developing rudiment of
fore wing; h., developing rudiment of hind wing; w., wing-pad. (After Graber;
twice natural size.)
 Development and Metamorphosis 43

development the young have to develop wings and make what other change
is necessary to reach the adult type, but the life is continually free and active
and the change is only a simple gradual transformation of the various parts
in which differences exist. A common locust is an excellent example of
an insect with such incomplete metamorphosis. Fig. 72 shows the develop-
ing locust at different successive ages, or stages, as these periods are called
because of their separation from each other by the phenomenon, common
to all insects, of moulting. As the insect grows it finds its increase of girth
and length restrained by the firm
inelastic external chitinized cuticle,
or exoskeleton. So at fixed periods
(varying with the various species
both in number and duration) this
cuticle is cast or moulted. From
a median longitudinal rent along
the dorsum of the thorax and head,
the insect, soft and dangerously
helpless, struggles out of the old
skin, enclosed in a new cuticle

which, however, requires some little

time to harden and assume its
proper colors (often protective).
F IG. 74. Metamorphosis, incomplete, of an
After each moulting young the
locust appears markedly larger and young after first moulting, showing
eggs; 5,
with ^ginning wing-pads; c, older stage with
its wing-pads better developed
r
complex wing-pads; D, adult with fully
(Fig. 73). But not until the final developed wings. (One-half larger than
in the case of the locust natural size.)
moulting
this is the fifth are the wings usable as organs of flight. So that there
is after all likely to be a rather marked difference between the habits of
the young and those of the adult of an insect with incomplete metamor-

phosis, that difference being primarily due to structural differences. The

young are confined to the ground, and their locomotion is limited to walking
or hopping. The adults can live, if they like, a life in the air, and they
have a means of locomotion of greatly extended capability.
The insects with complete metamorphosis are the beetles, the two-
winged flies, the butterflies and moths, the ichneumons, gall-flies, ants,
bees, and wasps, the fleas, the ant-lions, and several other small groups
of insects with less familiar names. In the case of all the thousands of
species in these groups, the young when hatched from the egg differ very
much in structure and appearance, and
and general economy,
also in habits
from the parents. Familiar examples of such young are the caterpillars
and "worms" of the moths and butterflies, the grubs of beetles, the mag-
44 Development and Metamorphosis

gots of the flesh- and house-flies, and the helpless soft white grubs in the
cells of bees and wasps. These strange young, so unlike their parents,
have the generic name larvae, and the stage or life of the insect passed as a
larva is known as the larval stage. In almost all cases these larvae have
mouth-parts fitted for biting and chewing, while most of the adults have
sucking- mouth parts; the larvae have only simple eyes and small inconspicu-

FIG. 75. Metamorphosis, complete, of monarch butterfly, Anosia plexippus. a, egg

(greatly magnified); b, caterpillar or larva; c, chrysalid or pupa; d, adult or imago.
(After Jordan and Kellogg. Natural size.)

ous antennae; the adults have both simple and compound eyes and well-
developed conspicuous antennae; the larvae may have no legs, or one pair or
two or any number up the adults have always three
to eight or ten pairs;

pairs; the larvae are wholly wingless, nor do external wing-pads (i.e.,

developing wings) appear outside the body during the larval stage; the
adults have usually two pairs (sometimes one or none) of fully developed

wings. Internally the differences are also great. The musculation of the
 Development and Metamorphosis 45
larva is like that of a worm, to accomplish wriggling, crawling, worm-like

locomotion; in the adult it is very different, particularly in head and thorax;

the alimentary canal is usually adapted in the larva for manipulating and

digesting solid foods; in the adult, usually (except with the beetles and
a few other groups), for liquid food; there may be large silk-glands in the
larva, which are rarely present in the
adult; the respiratory system of the larvae
of some flies and Neuroptera is adapted
for breathing under water; this is only
rarely true of the adults. The heart
and the nervous system show lesser dif-

ferences, but even here there is no iden-

tity the ventral nerve chain of the larvae
:

may contain twice as many distinct gan-

glia as in the adult.
Thelarva lives its particular kind of
life: grows and moults several times;
it

but externally it shows at no time any

more likeness to the adult than it did at FIG. 76. Larva, pupa, and adult of
the flesh-fly, Calliphora crythroce-
hatching. But after its last moult it
ap- phala, with complete metamor-
pears suddenly in the guise of a partially phosis. (Two times natural size.)
formed adult in (usually) quiescent mummy-like form, with the antennas,
legs, and wings of the adult folded compactly on the under side of the

body, and the only sign of life a feeble bending of the hind-body in re-
sponse to the stimulus of a touch. This is the insect of complete meta-
morphosis in its characteristic second stage (or third if the egg stage
is called first), the pupal stage. The
mummy is called pupa or chrysalid. As
the insect cannot, in this stage, fight or
run away from its enemies, its defence

lies in the instinctive care with which the

larva, just before pupation, has spun a
FIG. 77. Adult worker (a) and larva protecting silken cocoon about itself, or
(b) of honey-bee. (Adult natural has burrowed below the surface of the
size; larva twice natural size.)
ground, or has concealed itself in crack
or crevice. Ormay the fine harmonizing of the color and
the defence lie in

pattern of the naked exposed chrysalid with the bark or twig on which it
rests; it may be visible but indistinguishable. The insect as pupa takes
no food; but the insect as larva has provided for this. By its greed and
overeating it has laid up a reserve or food-store in the body which is drawn
on during the pupal stage and carries the insect through these days or weeks
or months of waiting for the final change, the transformation to the renewed
46 Development and Metamorphosis
active food-getting life of the adult or imaginal stage. Familiar examples
of this kind of metamorphosis, the real metamorphosis, are provided
by
the life monarch butterfly, the honey-bee, and the blow-fly. The great
of the
red-brown monarch lays its eggs on the leaves of a milkweed; from the
eggs
hatch in four days the tiny tiger-caterpillars (larvae) (Fig. 75) with biting
mouth-parts, simple eyes, short antenna;, and eight pairs of legs on its elon-
gate cylindrical wingless body. The caterpillars bite off and eat voraciously
bits of milkweed-leaf; they grow rapidly, moult four times, and at the end
of eleven days or longer hang themselves head downward from a stem or

FIG. 78. Brood-cells from honey-bee comb showing different stages in the metamor-
phosis of the honey-bee; worker brood at top and three queen-cells below; begin-
ning at right end of upper row of cells and going to left, note egg, young larva, old
larva, pupa, and adult ready to issue; of the large curving queen-cells, two are cut
open to show larva within. (After Benton; natural size.)

leafand pupate, i.e., moult again, appearing now not as caterpillars, but as
the beautiful green chrysalids dotted with gold and black spots. The form-
ing antennae legs and wings of the adult show faintly through the pupal
cuticle, but motionless and mummy-like each chrysalid hangs for about
twelve days, when through a rent in the cuticle issues the splendid butterfly
with its coiled-up sucking proboscis, its compound eyes, long antennae, its
three pairs of slender legs (the foremost pair rudimentary), and its four great
red-brown wings. The queen honey-bee lays her eggs, one in each of the
scores of hexagonal cells of the brood-comb (Fig. 78). From the egg there
hatches in three days a tiny footless, helpless white grub, with biting mouth-
parts and a pair of tiny simple eyes. The nurses come and feed this larva
steadily for five days; then put a mass of food by it and "cap" the cell; the
larva has grown by this time so as nearly to fill the cell. It uses up the

stored food, and "changes" to the pupa, with the incomplete lineaments
of the adult bee. It takes no more food, but lies like a sleeping prisoner
 Development and Metamorphosis 47
in closed cell for thirteen days, and then it awakens to active life, gnaws
its

itsway through the cell-cap and issues into the hive-space a definitive honey-
bee with all the wonderful special structures that make the honey-bee body
such an effective little insectean machine. The blow-fly (Fig. 76) lays a hun-
dred or more little white eggs on exposed meat. From these eggs come in
twenty or thirty hours the tiny white wriggling larvae (maggots), footless, eye-
less, wingless, nearly headless, with a single pair of curious extensile hooks
for mouth-parts. For ten to fourteen days these larvae squirm and feed and
grow, moulting twice in this time; they then pupate inside of the larval
cuticle, which becomes thicker, firmer, and brown, so as to enclose the deli-
cate pupa in a stout protective shell. The blow-fly now looks like a small
thick spindle-shaped seed or bean, and this stage lasts for twelve or fourteen

ftmth
. Amur

FIG. 79. Dipterous larvae showing (through skin) the imaginal discs or buds of wings,
these buds being just inside the skin. A, larva of black fly, Simulium sp.; B, anteiior
end of larva of midge, Chironomus sp. C, anterior end, cut open, of larva of giant
;

crane-fly, Holorusia rubiginosa; h.pr., bud of prothoracic respiratory tube; h.pl.,

bud of prothoracic leg; h.m-w., bud of mesothoracic wing; h.ml., bud of mesothoracic
leg; h.mtb., bud of metathoracic balancer; h.mtl., bud of metathoracic leg. (Much
enlarged.)

days. the winged imago, the buzzing blow-fly, as we best know it,
Then
breaks itsout. In the house-fly the same kind of life-history, with
way
complete metamorphosis of the extremest type, is completed in ten days.
Nor do we realize how really extreme and extraordinary this metamorpho-
sis is until we study the changes which take place inside the body, as well
as those superficial ones we have already noted.
The natural question occurs to the thoughtful reader: "Is the meta-
morphosis or transformation in the postembryonal development of such
insects as the butterfly, bee, and blow-fly as sudden or discontinuous and
" The
as radical as the superficial phenomena indicate? answer is no, and
yes; the metamorphosis is not so discontinuous or saltatory and yet is
even more radical and fundamental than the external changes suggest. To
48 Development and Metamorphosis
take a single example, the case of the blow-fly (admittedly an extreme
one),
the phenomena of internal change are, put briefly, as follows: The imaginal
wings, legs, and head-parts begin to develop as deeply invaginated little
buds of the cell-layer of the larval skin early in larval life. This develop-
ment is gradual and continuous until pupation, when the wing and leg rudi-

jam.
FIG. 80. Stages in development of wing-buds in the larva of the giant crane-fly,
Holorusia rubiginosa (the wing-buds have been dissected out and sectioned, so
as to show their intimate anatomy). A, B, C, D, four stages successively older ch.,
chitinized cuticle; hyp., hypoderm or cellular layer of skin; tr., trachea; trl.,
tracheoles; p.m., peritrophic membrane; w., developing wing; t.v., tracheal branch
indicating position of future wing-vein. (Greatly magnified.)

ments and the new head are pulled out upon the exterior of the body. Just
before pupation, when the larva has given up its locomotion and feeding,
the larval muscles, tracheae, salivary glands, alimentary canal, and some other
and rapidly break wholly down, so that in the
tissues begin to disintegrate,

pupa there
appear be no
to internal organs except the nervous system,

reproductive glands, and perhaps the heart, but the whole interior of the
 Development and Metamorphosis 49

body is filled with a thick fluid in which float bits of degenerating larval
tissue. At the same time withthis radical histolysis or breaking down of

tissue a rapid histogenesis or developing of imaginal parts from certain

groups of undifferentiated primitive cells, derived probably mostly from
the larval skin-cells, is going on. Thus many of the larval organs and tissues,
instead of going over into the corresponding imaginal ones, wholly disinte-

grate and disappear, and the imaginal parts are newly and independently
derived. In connection with the
breaking down of the larval tissues

phagocytes or freely moving, tissue-

eating, amosboid blood-cells play an
important part, although one not

yet fully understood. They are

either the causal agents of the

histolysis, or are assisting agents in

it,the tissue disintegration beginning

independently, or a recent sugges-

tion they are perhaps more truly
to be looked on as trophocytes,
that is,
carriers of food, namely,
disintegrating tissue, to the develop- FIG. 81. A cross section of the body of the
pupa of a honey-bee, showing the body-cavity
ing centers of the imaginal parts. filled with disintegrated tissues and
phago-
Much investigation remains to be cytes, and (at the bottom) a budding pair
of of the the larvae
done on this interesting subject legs adult, being
wholly legless. Photomicrograph by George
of histolysis and histogenesis in O. Mitchell; greatly magnified.)
insects with complete metamor-
phosis, but enough has been already accomplished to show the basic and
extreme character of the transformation from larva to adult.
If we ask for the meaning of such unusual and radical
changes in the
development of insects, we confront at once an important biological prob-
lem. Most biologists believe that in a large and general way the develop-
ment of animals is a swift and condensed recapitulation of their evolution;

meaning by development the life-history or ontogeny of an individual, and

by evolution the ancestral history or phylogeny of the species. According
to this "biogenetic law" the interpretation of the significance of the various

stages and characters assumed by an animal in the course of its development

from single fertilized egg-cell to the complex many-celled definitive adult
stage is simple: These stages correspond to various ancestral ones in the
long genealogical history of the species. Every vertebrate, for example, is
at some period in its development more likea fish than any other living
kind of animal ;
it has
gill-slits in its throat, is tailed, and is indeed a fish-
like creature. This is its particular developmental stage, corresponding
5 Development and Metamorphosis
to the ancestral fish-like ancestors of all vertebrates. Do then the larvae
and pupae of insects with complete metamorphosis represent ancestral stages
in insect evolutionary history? In some degree the larval stage does, but
in no degree does the pupal.
Insects are certainly not de-
scended from an animal that,
like a pupa, could neither move

FIG. 82. A bit of degenerate muscle from tussock- nor eat and which had no in '
moth, Hemerocampa leucostigma. Note phago- ternal organs except a nervous

system, heart, and rudimentary

reproductive glands. Biologists
recognize that the exigencies of life during adolescence may profoundly
modify what might be termed the normal course of development. As
long as the developing animal is shielded from the struggle for existence,
is provided with a store of food and protected from enemies
by lying in an
egg-shell or in the body of the mother, it may pursue fairly steadily its reca-
pitulatory course of development; but once emerged and forced to shift for

FIG. 83. Degenerating muscle from pupa of giant crane-fly, Holorusia rubiginosa, show-
ing phagocytic cells penetrating and disintegrating the muscle-tissue. (Greatly
magnified.)

itself, it must be, at whatever tender age it is turned out, or whatever ancient
ancestor it is in stage of simulating, adapted to live successfully under the

present-day and immediate conditions of life. If the butterfly gets hatched

long before it has reached its definitive butterfly stage, and while it is in
a stage roughly corresponding to some worm-like ancestors and from such
ancestors insects have undoubtedly descended it must be fitted to live
 Development and Metamorphosis

successfully a crawling, squirming, worm-like life. That those insects which

hatch as worm-like larvae do in fact owe their wingless, worm-like body con-
dition partly to being born in a stage simulating a worm-like ancestor is proba-

FIG. 84. Degeneration, without phagocytosis, of salivary glands in old larva of giant
crane-fly,Holorusia rubiginosa. A, cross-section of salivary gland before degen-
eration has begun; B, cross-section of salivary gland after degeneration has set in.
(Greatly magnified.)

bly true. But to be a successful worm demands very different bodily adapta-
tions from those of a successful butterfly. And so far does the larval butterfly
go, or so far has it been carried, in meeting these demands that nature finds it

more economical to get into figurative language

or easier to break down almost wholly the larval

body after a new food-supply for further develop-

ment has been got and stored away, and to
build up from primitive undifferentiated cell begin-
nings the final definitive butterfly body, than to
make over these very unlike larval parts into the
adult ones. The pupal stage, quiescent, non-food
taking, and defended by a thick chitinous wall,
often enclosed in a silken cocoon, buried in the

ground or crevice, or harmonizing so perfectly with

itsenvironment as to be indistinguishable from it,
is the chief period of this radical and marvelous FIG. 85. Cross-section
of newly developing
breaking down and building anew. It is an inter- muscle in pupa of
polated stage in the development of the butterfly honey-bee, Apis mel-
lifica. (Greatly mag-
corresponding to nothing in the phyletic history;
nified.)
an adaptation to meet the necessities of its life-
conditions. To my mind, this is the interpretation of the phenomena of

complete metamorphosis.
 CHAPTER III

THE CLASSIFICATION OF INSECTS

As has been explained in the preceding chapter, insects are primarily classi-
fied on the basis of their postembryonic development. Insects with incom-

plete metamorphosis, that is, those which do not undergo a non-feeding,

usually quiescent, pupal stage in their development are believed to be more

nearly related to each other than to any of the insects which undergo a so-
called complete metamorphosis. So they are spoken of collectively as the
Hemimetabola, while all the insects with a distinct pupal stage are called
the Holometabola. But when one has collected an adult insect, as a fly
or moth or grasshopper, and wishes to classify it, this primary classification
based on character of development often cannot be made for lack of informa-
tion regarding the life-history of the particular insect in hand. The next
grouping is into orders, and this grouping is based chiefly on structural
characters, and corresponds to one's already more or less familiar knowledge
of insect classification. Thus the beetles with their horny fore wings
all

constitute one order, the Coleoptera; the moths and butterflies with their
scale-covered wings another order, the Lepidoptera; the two- winged flies
the order Diptera, the ants, bees, wasps, and four-winged parasitic flies
the order Hymenoptera, and so on. So that the first step in a beginner's
attempt to classify his collected insects is to refer them to their proper orders.
Now while entomologists are mostly agreed w!th regard to the make-up
of the larger and best represented orders, that is, those orders containing
the more abundant and familiar insects, there are certain usually small,
obscure, strangely formed and more or less imperfectly known insects with
regard to whose ordinal classification the agreement is not so uniform. While
some entomologists incline to look on them simply as modified and aberrant
members of the various large and familiar orders, others prefer to indicate
the structural differences and the classific importance of these differences
by establishing new orders for each of these small aberrant groups. Most
entomologists of the present incline toward this latter position, so that whereas
Linnaeus, the first great classifier of animals, divided all insects into but
seven orders, the principal modern American * text-book of systematic ento-

* Comstock,
J. H., A Manual of Insects, 1898.
52
 The Classification of Insects
53

mology recognizes nineteen distinct ones. This does not mean, of course, that
twelve new
orders of insects have been found since Linnaeus's time, although
two or three of the orders are in fact founded on insects unknown to him,
but means that certain small groups classified by Linnaeus simply as families
in his large orders have been given the rank of distinct orders by modern

systematists. And as our knowledge of insects and their relationship to

each other is certainly much larger now than it was one hundred and fifty
years ago, we may feel confident that the many-order system of classifica-

tion more nearly a true expression of the natural interrelationships of

is

insects than was the old seven-order system. But not all entomologists
agree on the nineteen-order system. Few, indeed, still use the Linnsean
system, but many believe that the division of the insect class into nineteen
orders gives too much importance to certain very small groups and to some
others which are not markedly aberrant, and these entomologists recognize
a lesser number of orders, varying with different authors from nine to about
a dozen. In this book we shall adopt the nineteen-order system as used
in Comstock's Manual. In the first place the author believes that this classi-
fication best represents our present knowledge of insect taxonomy; in the

second place this is the classification taught by nearly all the teachers of

entomology in America.
To determine the order to which an insect belongs we make use of a
classifying table or key. In the Key to Orders which follows this para-
graph, all the insect orders are characterized by means of brief statements of
structural features more or less readily recognized by simple inspection of
the superficies of the body; to determine some of the conditions a simple
lens or hand-magnifier will be needed. The orders are so arranged in the
key that by choosing among two or more contrasting statements the student
may "trace" his specimen to its proper order. Inspection of the Key with
an attempt or two at tracing some familiar insect, as a house-fly, moth, or
wasp whose order is already known, will make the method of use apparent.
It must be borne in mind that young insects, such as caterpillars of moths,

grubs of beetles, and the wingless nymphs of locusts, dragon-flies, etc., cannot
be classified by this key. Indeed the young stages of most of the insects
which we know well as adults are unknown to us, and there is, besides, such
manifold adaptive variety in the external structure of those forms which we
do know that no key for the classification into orders of immature insects
can now be made.
54 The Classification of Insects

KEY TO THE ORDERS OF INSECTS.

(ARRANGED BY PROF. H. E. SUMMERS.)

(For adult insects only. If in any paragraph all the italicized characters agree with
the specimen in hand, the remaining characters need not be read; these latter are for use
in doubtful cases, or where the organs characterized in italics are rudimentary or absent.
The technical terms used in this Key have all been denned in Chapter I.)

A. Primitive wingless insects; mouth-parts well developed, but all except the apices of the
mandibles and maxilla -withdrawn into a cavity in the head; tarsi (feet) always one-
or two-clawed; body sometimes centiped-like, with well-developed abdominal legs,
in this case tarsi two-clawed (The simplest insects.) APTERA.
AA. Normally -winged insects, wings sometimes rudimentary or absent; mouth-parts
not withdrawn into a cavity in the head.
B. Mouth-parts, when developed, with both mandibles and maxilla fitted for biting;
abdomen broadly joined to thorax; tarsi never bladder-shaped; when mouth-
parts are rudimentary, if the wings are two, there are no halteres (p. 303) ; if
the wings are four or absent, the body is not densely clothed with scales.
C. Posterior end of abdomen -with a pair of prominent unjointed forceps-like

appendages; fore wings, when present, short, veinless, horny or leathery.

(Earwigs.) EUPLEXOPTERA.
CC. Posterior end of abdomen usually without prominent unjointed forceps-like
appendages; when these are present the fore wings are always developed,
veined.
D. Fore wings, when present, veined and membranous, parchment-like or
leathery; when absent, the labium (under-lip) either cleft in the
middle, or the mouth-parts prolonged into a distinct beak.
E. Fore wings, when present, thicker than hind wings, somewhat
leathery or parchment-like; hind wings folded several times
lengthwise, like a fan, in repose; when wings are absent, pro-
thorax large.
(Locusts, crickets, cockroaches, etc.) ORTHOPTERA.
EE. Fore wings membranous, of same structure as hind wings;
hind wings usually not folded, but occasionally folded like a fan;
when wings are absent, prothorax small.
F. Antenna inconspicuous.
G. Hind wings smaller than fore or absent; posterior end of
abdomen with two or three many-jointed filaments.

(May-flies.) EPHEMERIDA.
GG. Hind wings not smaller than fore; postejior end of
abdomen without many-jointed filaments.
(Dragon-flies and damsel-flies.) ODONATA.
FF. Antenna conspicuous.
G. Tarsi less than five-jointed; labium cleft in the
middle.
H. Wings always present, although sometimes very
small; hind wings broader than fore wings,
folded in repose; prothorax large, nearly flat
on dorsal surface.
(Stone-flies.) PLECOPTERA.
 The Classification of Insects 55
HH. Hind wings, when present, not broader than fore
wings, not folded in repose; prothorax small,
collar-like.
I. Tarsi four-jointed; wings, when present,
equal in size (Termites.) ISOPTERA.
II. Tarsi one- to three-jointed.
J. Tarsi one- or two-jointed; always
wingless.
(Biting bird-lice.) MALLOPHAGA.
JJ. Tarsi usually three-jointed; occasionally
two-jointed, in which case wings always
present, fore wings larger than hind
wings. (Book-lice, etc.) CORRODENTIA.
GG. Tarsi five-jointed, but with one joint sometimes
difficult to distinguish; labium usually entire in
middle, sometimes slightly emarginate.
H. Wings, when present, naked or slightly hairy;
hind wings with or without folded anal space;
in former case prothorax large and nearly
flaton dorsal surface; in wingless forms
mouth prolonged into a distinct beak.
I. Mouth-parts not prolonged into a distinct
beak, at most slightly conical.
(Dobsons, ant-lions, etc.) NEUROPTERA.
II. Mouth-parts prolonged into a distinct beak.
(Scorpion-flies, etc.) MECOPTERA.
HH. Wings, when present, thickly covered with hairs;
hind wings usually with folded anal space; pro-
thorax small, collar-like; mouth not prolonged
intoa beak. (Caddis-flies.) TRICHOPTERA.
DD. Fore wings, when present, veinless; horny or leathery; when absent,
labium entire, and mouth-parts not prolonged into a distinct beak.
(Beetles.) COLEOPTERA.
BB. Mouth-parts, when developed, more or less fitted for sucking; sometimes also
fitted in part (the mandibles) for biting: in this case either (i) base of abdomen

usually strongly constricted, joined to thorax by a narrow peduncle, or (2) the

tarsi bladder-shaped, without claws; when mouth is rudimentary either the

wings are two and halteres are present, or the wings are four or none and
the body (and wings if present) are densely clothed with scales.
C. Prothorax free; body (and wings if present) never densely clothed with
scales; maxillary palpi usually absent; when present, tarsi bladder-
shaped, without claws.
D. Tarsi bladder-shaped, without claws; wings four (sometimes absent),
narrow, fringed with long hairs; maxillae triangular, with palpi.
(Thrips.) THYSANOPTERA.
DD, Tarsi not bladder-shaped, usually clawed; wings not fringed with
long hairs; maxilla (when mouth is developed) bristle-like, without
palpi. (Bugs.) HEMIPTERA.
CC. Prothorax not free;maxillary palpi present, sometimes rudimentary
and difficult to see, in which case body (and wings if present) densely
clothed with scales; tarsi never bladder-shaped, usually clawed.
56 The Classification of Insects

D. Mandibles often rudimentary, when present bristle-like.

E. Wings (sometimes wanting), clothed with scales; body

four
covered thickly with scales or hairs; mouth, when developed, a
slender sucking proboscis, closely coiled under head.
(Moths and butterflies.) LEPIDOPTERA.
EE. Wings two (or wanting), naked or with scattered hairs; hind
wings in winged jorms represented by halteres; body either
naked or with scattering hairs; mouth a soft or horny beak, not
coiled under head.
F. Prothorax poorly developed, scarcely visible from dorsal
side (Flies.) DIPTERA.
FF. Prothorax well developed, distinctly visible from dorsal
side; wings never present (Fleas.) SIPHONAPTERA.
DD. Mandibles well developed, fitted /or biting; wings four (sometimes
two or none), naked or with scattered hairs.
(Ichneumon-flies, gall-flies, wasps, bees, and ants.) HYMENOPTERA.

After one has classified an insect in its proper order there remains, first,
the determination of the family (each order being composed of from one
to many families), then of the genus (each family comprising one to many

genera), and finally of the particular species of the genus (each genus includ-
ing one to many species). This ultimate classification to species, however,
will be possible to the amateur in comparatively There are so
few cases.
many species of insects (about 300,000 are known) that would require it

many shelves of books to contain the descriptions of them all. As a matter

of fact, in only a few orders have the descriptions of the species been brought

together in manuals available for general students. For the most part the
descriptions are scattered in scientific journals printed in various languages
and wholly inaccessible to the amateur. There are less than 1000 different
species of birds in North America; there are more than 10,000 known
species of beetles. Now when one recalls the size of the systematic man-
uals of North American birds, and realizes that ten such volumes would
include only the insects of one order, it is apparent that complete manuals
of North American insects are out of the question. Except in the case of
the most familiar, wide-spread, and readily recognizable insect species we
must content ourselves with learning the genus, or the family, or with the
more obscure, slightly marked, and difficult members of certain large groups,
as the beetles and moths, simply the order of our insect specimens.
When one has determined the order of an insect by means of the above
key he should turn to the account of this particular order in the book (see
index for page) and find the keys and aids to the further classification of
the specimen which the author has thought could be used by the general
student. Comparison with the figures and brief descriptions of particular
species which are given in each order may enable the amateur to identify
the exact species of some of his specimens. But the specific determination
 The Classification of Insects 57
of most of the insects in an amateur's cabinet (or in a professional ento-
mologist's either, for that matter) will have to be done by systematic
specialists in the various insect groups. Few professional entomologists
undertake to classify their specimens to species in more than the one or
two orders which they make their special study. Duplicate specimens should
be given numbers corresponding to those on specimens kept in the cabinet,
and be sent to specialists for naming. Such specialists, whose names can
be learned from any professional entomologist, have the privilege of retain-
ing for their own collections any of the specimens sent them.
 CHAPTER IV

THE SIMPLEST INSECTS (Order Aptera)

ERTAIN household pests which are

not moths and do not look like

fish, but which are com ^L^J rnonly called "fish-moths" (Fig. 86), are
our most familiar repre sentatives of the order of "simplest in-
sects." The "fish" part of the name comes from the
covering of minute scales which gives the body a silvery
appearance, and the "moth" part is derived from our
habit of calling most household insect pests "moths."
Thus we speak of "buffalo-moths" when we refer to the

carpet-feeding hairy larvae of certain beetles. When we

say clothes-moths we are really using the word moth
accurately, for in their adult condition these pests are
true moths, although the injury to clothing is wholly done

by the moth in its young or caterpillar stage. FIG. 86. The fish-
Besides the fish-moths other not unfamiliar Aptera are moth, Lepisma
" " saccharina. (After
the tiny springtails (Fig. 87), which sometimes occur Howard and Mar-
in large numbers on the surface of pools of water or on latt: twice natural

snow in the spring. Others may be easily found in damp size.)

decaying vegetable matter, as discarded straw or old toadstools. They are

provided with an odd spring on the under side of the body by means
little

of which they can leap from a few inches to a foot

or more into the air. Hence their common name.
In the order Aptera are included the simplest of
living insects. By "simplest" is meant most primi-
tive, most nearly related to the ancestors of the whole

FIG. 87. The

insect class. Also, as might be expected, these most
pond-sur-
face springtail, Smyn- primitive insects are simplest in point of bodily struc-
thurus aquatic 11 s. ture; but in this respect they are nearly approached
(After Schott; much
enlarged.) by simple-bodied members of several other orders.
These latter forms, however, have a simple body-
structure due to the degradation or degeneration of a more complex type.
58
 The Simplest Insects 59

It is familiarknowledge that animals which live parasitically on others, or

which adopt, a very sedentary life, show a marked degeneration of body
structure, an acquired simplicity due to the loss of certain parts, such as
organs of locomotion (wings, legs), and of
orientation (eyes, ears, feelers, etc.). Thus
the parasitic biting bird-lice (order Mal-
lophaga, see p. 113), which live their whole
lives through on the bodies of birds, feeding
on the feathers, are all wingless and of gener-

ally simple superficial structure. They are

nearly as simple externally perhaps as the

Aptera, but we believe that they are the
degenerate descendants of winged and in
other ways more complexly formed ancestors.

Similarly certain species of in insects

nearly all orders have adopted a life-habit

which renders flight unnecessary, and these tion of the ovarial tubes in three
Apteran genera. A, Japyx; B,
insects having lost their wings are in this
Lepisma; C, Campodea. (After
character simpler than the winged kinds. Targioni-Tozzetti; much en-
Examples of such insects are the worker larged.)

ants and worker termites, many household insects, as the bedbugs and fleas,
and many ground-haunting forms, as some
of the crickets, cockroaches, and beetles.
The Aptera, however, owe their sim-
plicity to genuine primitiveness; among all

living insects they are the nearest repre-

sentatives of the insectean ancestors. But
not all the Aptera are "simplest." That
is, within the limits of this small order a
considerable complexity or specialization of
structure is attained, although all the

Aptera are primitively wingless, as the

name of the order indicates.
These insects develop "without meta-
"
morphosis is, the young (Figs. 90
;
that
and 94) are almost exactly like the parents
FIG. 89. Diagrammatic figures show- except
in size. They have simply Jo grow
ing the respiratory system in three larger and to become mature. In internal
Apteran genera. A, Machilis; B, ,1-1
structure the simpler Aptera show some
Nkoletia; C, Japyx. (After Tar-
gioni-Tozzetti; much enlarged.) most interesting conditions. Their internal
systems of organs have a segmental character corresponding to the external
segmentation of the body. The ovarial tubes, which are gathered into
60 The Simplest Insects

two groups or masses, one on each side of the body, in all other insects
(Fig. 66), are separate and arranged segmentally in Japyx (Fig. 88), and
less markedly so in Machilis; the respiratory system of Machilis (Fig. 89)
consists of nine pairs of distinct, segmentally arranged groups of tracheae

(air-tubes), while the ventral nerve-cord has a ganglion in almost every seg-
ment of the body. As insects are certainly descended from ancestors whose
bodies were composed of segments much less interdependent and coordi-
nated than those of the average living insect, those present-day insects which
have the body both externally and internally most strongly segmented are
believed to be the most generalized or primitive of living forms. In addi-
tion to the segmented character of the internal organs we have also another

strong evidence of the primitiveness of the order in the possession by several

Aptera of rudimentary but distinct external pairs of appendages on the
abdominal segments, appendages undoubtedly homologous with the thoracic
legs, and probably well developed in the insect ancestors as abdominal legs
like those of the centipeds.
The order Aptera is composed of two suborders, which may be dis-

tinguished as follows:
Abdomen elongate, composed of ten segments, and bearing long bristle-like or
shorter forceps-like appendages at its tip; no sucker on ventral side of first
abdominal segment; antennae many-segmented THYSANURA.
Abdomen short and robust, composed of six segments, and usually with a forked
spring at tip (usually folded underneath the body), and with a ventral sucker
on first abdominal segment; antennas 4- to 8-segmented COLLEMBOLA.

THYSANURA. This suborder includes three families (a problematical

fourth family is found in Europe), as follows:
Body covered with scales LEPISMID^E
Body not covered with scales.
Tip of abdomen with forceps-like appendages JAPYGID^E.
Tip of abdomen with slender many-segmented appendages CAMPODEID.E.

To the last family in the above key belongs the interesting creature

Campodea staphylinus (Fig. 90) which zoologists regard as the most primi-
,

tive living insect. It is small, white, flattened, wingless, and so soft-bodied

and delicate that can hardly be picked up uninjured with the most deli-
it

cate forceps. It is about \ inch long (exclusive of caudal appendages) and ,

is to be looked for under stones and bits of wood. I have found it in Ger-

many, in New York, and in California, which indicates its wide distribu-
tion. Other collectors have taken it in Italy, England, and in the Pyrenees.
It is said to live also inEast India. Is it not a little surprising that this
most primitive, wholly defenceless, and ancient insect should be able to live
successfully the world over in the face of, and presumably in competition
with, thousands of highly developed specialized modern insect forms? It
 The Simplest Insects 61

is a striking proof that Nature does not inevitably crush out all of her
first trials in favor of her later results!
The Campodeidae contain another
genus, Nicoletia (Fig. 91), one species of
which, N. lexensis, has been found in Cali-
fornia and Texas, and which may be dis- /
*
tinguished from Campodea by its posses-
ft

sion of three caudal appendages instead

of two as in the latter form.
The Japygidae include but a single
genus, Japyx, represented countryin this

by two described species and several as yet

undescribed forms found at Stanford Uni-
versity. Japyx subterraneus is a species
first found under stones at the mouth of
a small grotto near the Mammoth Cave
(Kentucky). Japyx (Fig. 92) is larger FlG 9- -
Young and adult of Cam-
than Campodea, being about one-half inch
long, and is readily recognized by its caudal size indicated by line.)

forceps. Like Campodea its body is white and soft.

The Lepismidae include the familiar household fish-
moths and a number of similar forms which live under
stones and logs in soft soil at the bases of tree-trunks,
under dead leaves in woods, and sometimes on the damp
sand of seashores. Three genera of this family occur
in North America, which may be distinguished as
follows :

Caudal appendages short; prothorax very wide and body

behind it tapering rapidly LEPISMINA.
Caudal appendages long body elongate and tapering
;

gradually backward.
Eyes large and close together MACHILIS.
Eyes small and far apart LEPISMA.

Lepisma is best known by the species L. saccharina

(Fig. 86), which is the silverfish or fish-moth of the
house. It is silvery white, with a yellowish tinge on

the antennae and legs, and is from one-third to two-

fifths f an inch n g' The three lon g Caudal a PP en -
l
FIG. gi.-Nicoletiate X-
ensis, from Califor- dages, characteristic of the genus, are conspicuous. It
nia. (Eight times nat-
f ee<^ s chiefly on sweet or starchy materials, sometimes
ural size.) . ..... .

doing much damage in libraries, where it attacks the

bindings. It attacks starched clothing, eats the paste off the wall-paper,
 The Simplest Insects

causing it to loosen, and infests dry starchy foods. It runs swiftly and
avoids the light. It can be fought by sprinkling fresh

pyrethrum powder in bookcases, wardrobes, and

pantries. Another species, L. domestica (Fig. 93),
called the bake-house silverfish, is often common
about fireplaces and ovens, running over the hot
metal and bricks with surprising immunity from the
effects of the heat. This habit has gained for it in
England, according to Marlatt, the name of "fire-
brat." It can be distinguished from the species

saccharina by the presence of dark markings on the

FIG. 92. Japyx sp., from back. Both saccharina and domestica are common
.California. (Five times in
natural size.)
England, and saccharina probably came to this
'

country from there.

Machilis (Fig. 95) does not occur in houses, but is more common than
Lepisma outdoors. It is to be found under stones, in the soil around the
base of tree-trunks, among dead leaves and fallen pine-needles, and at least
one species occurs in the sand of sea-beaches.

FIG. 93. FIG. 94.

FIG. 93. Thefish-moth, Lepisma domestica. (After Howard and Marlatt; a little

larger than natural size.)

FIG. 94. Young and adult of Lepisma sp., from California. (Twice natural size.)

COLLEMBOLA. The springtails, mostly of microscopic size, and wholly

unfamiliar to any but persistent explorers of nature, comprise many more
species than the Thysanura. Their most distinctive character is the pos-
session, by most of them, of the forked spring (Figs. 96 and 97), by
means of which they leap vigorously when disturbed. This spring is
 The Simplest Insects

attached to the next to last body segment or to the antepenultimate one.

It consists of a basal part and of two terminal processes.

It is carried bent forward under the body, with the bipartite

tip held in a little catch on the third abdominal segment.

In some species the catch is lacking. The springtails also
possess a curious organ on the ventral aspect of the first
abdominal segment which appears to be a small projecting
sucker or tube. This sucker is often more or less divided
into two parts, in one family consisting plainly of two

elongate, delicate tubes (Figs. 96 and 97). The use of

this peculiar structure has not been definitely determined.

Some entomologists think that it serves as a clinging organ,

enabling the insect to attach its body firmly to the object
upon which it rests. Others believe that the sucker serves
in some way to take up moisture, while still others be-
lieve it to aid in respiration. The Collembola as well
as the Thysanura cannot live in a dry atmosphere.
This suborder is divided into five families, as follows
(MacGillivray) :

A. Spring wanting .APHORURID.E.

FIG. 95. Machi-
AA. Spring present. lis sp.,from Cali-
B. Spring arising from ventral side of fornia. (Three
antepenultimate abdominal segment. times natural
PODURID.E. size.)

BB. Spring arising from ventral side of penultimate abdom-

inal segment.
C. Abdomen elongate, cylindrical, much longer than
FIG. 9 6.-The spotted
broad ENTOMOBRYID*.
springtail, Papirius
CC. Abdomen globular, but little larger than broad.
maculosus, with spring D. Terminal segment of antennae long, ringed,
folded underneath SMYNTHURID^:.
body (Natural DD Terminal seg ment of the antennae short, with
length, 2 mm.) .
,
. .
a whorl of hairs PAPIRIID^E.

Of these five families the members of one, the Aphoruridae, in which

the spring is wanting, are non-saltatorial. In all of
the others leaping is a characteristic habit. The
Smynthuridae and the Papiriidaa are represented by
but one genus each, viz., Smynthurus and Papirius.
Smynthurus hortensis is a common form in gardens,
and may be called the "garden-flea." It is found
in the Eastern States in and June "upon the
May FIG. 97. The spotted
leaves of young cabbage, turnip, cucumber, and springtail, Papinus mo fu-
losus, with spring extended,
various other plants, and also on the ground. It (Natural length, 2 mm.)
 The Simplest Insects

is dull black, with head, legs, and bases of the antennae rust-color."
Smyn-
thurus aquaticus (Fig. 87) often occurs in great numbers on the surface of
pools. The insects look like tiny black spots on the water surface, but a
little observation soon reveals their
lively character.
The Poduridae and Entomobryidae
are represented in North America by
twelve and fourteen genera respec-
I It IS' nfl'OT xft
tively '
Many of the Podurids are
il ff '
il u'fJf US' covered with scales and are often
If H\ 1 W/JB / ^fiS
prettily colored and patterned. The
scales (Fig. 98) are very minute and
bear many fine lines and cross-lines,
regularly arranged. On this account
FIG. 98. FIG. 99. t ne y are much used as test
objects
FIG 9 8.-Scales from a springtail. (After for
microscopes
r the qualit
* of the
Murray; greatly magnified.) _ >
FIG. 99. The snow-flea, Achorutes nivicola. lens being determined by its capacity
(After Folsom; much enlarged.) to reveal their extremely fine mark-

ings. One of the most interesting Podurids is

the snow-flea, Achorutes nivicola (Fig. 99), which

gathers in large numbers on the surface of snow
in the late spring. Comstock says that the
snow-flea is sometimes a pest where maple-
sugar is made, the insects collecting in large
quantities in the sap.
An interesting representative of the Entomo-
bryidae the house springtail, Lepidocyrtus ameri-
is

canus (Fig. 100), said by Marlatt to be "not

infrequently found in dwellings in Washington."
FIG. ioo. The American
It is about one-tenth of an inch long, silvery
springtail, Lepidocyrtus
gray, with purple or violet markings. In Europe americanus, ventral aspect,
showing spring folded un-
also one species of springtail is common in derneath body. (After
houses. As these insects live on decaying vege- Howard and Marlatt ;
table matter, they probably do no special harm enlarged.) much
in the house. They especially frequent rather moist places, and may often
be found in window-plant boxes and conservatories.
 CHAPTER V
THE MAY-FLIES (Order Ephemerida) and STONE-
f ' - FLIES (Order Plecoptera)

AY-FLIES, lake-flies, or shad-flies, common names for

the insects of the order Ephemerida, are familiar to

people who live on the shores of lakes or large rivers,

but are among the unknown insects to most high-and-
dry dwellers.
Travelling down the St. Lawrence River from
Lake Ontario to Quebec one summer, I had hosts of
day-long companions in little May-flies that clung to
my clothing or walked totteringly across my open book. The summer
residents of the Thousand Islands get tired of this too-constant com-

panionship, and look resentfully on the feeble shad-fly as an insect pest.

One evening in August, 1897, mv attention, with that of other strollers along
the shore promenade at Lucerne, was called to a dense, whirling, tossing
haze about a large arc light suspended in front of the great Schweizerhof.
Scores of thousands of May-flies, just issued from the still lake, were in
violent circling flight about the blinding light, while other thousands were

steadily dropping, dying or dead, from the dancing swarm to the ground.
Similar sights are familiar in summer-time in this country about the lights
of bridges, or lake piers and shore roads. This flying dance is the most
conspicuous event in the life of the fully developed, winged May-fly, and
indeed makes up nearly all of it. With most species of May-flies the winged
adult lives but a few hours. In the early twilight the young May-fly floats
from the bottom of the lake to the surface, or crawls up on the bank, the
skin splits, the fly comes forth full-fledged, joins its thousands of issuing

companions, whirls and dances, mates, drops its masses of eggs on to the
the lake's surface, and soon flutters and falls after the eggs. It takes no

food, and dies without seeing a sunrise. Sometimes the winds carry dense
clouds of May-flies inland, and their bodies are scattered through the streets
of lakeside villages, or in the fields and woods. Sometimes the great swarms
65
66 The May-flies and Stone-flies

fall to the water's surface and there are swept along by wind and wave,
up in thick winrows, miles long,
until finally cast
on the lake beach. Millions of dead May-flies
are thus piled up on the shores of the Great
Lakes.
We call the May-flies the Ephemerida, after

the Ephemerides of Grecian mythology, and the

name truly expresses their brief existence above
water. But they have lived for a year at least
before this, or for two or even three years, as

wingless, aquatic creatures, clinging concealed

to the under side of stones in the lake or stream
bottom, or actively crawling about after their food,
which consists of minute aquatic plants and animals
or bits of dead organic matter. In this stage their

whole environment, habits, and general appearance are

radically different from those of the brief adult life. We
can only guess, if our curiosity compels us to attempt some
explanation, at the manner and the cause of such a

strange life-history. What advantage is there in such a

specialized condition that Nature could not have arrived
at by less indirect means ? What is indeed the utility of
the whole modification? The quick answer "utility,"
which is such strange structural and
to account for all

physiological conditions on the basis of useful adapta-

tions brought about by the slow but persistent action
of natural selection, leaves us, confessedly, answered
simply on a basis of belief. In hundreds of cases that
may come under our observation, in how few are we
really able to perceive a reason-satisfying course of adap-
tive development based on the selection of useful small
fluctuating variations?
The eggs of the May-fly fall from the body of the
mother to the water's surface in two packets, which,
however, break up while sinking, so that the released

FIG. 101. May-flies about an electric lamp.

 The May-flies and Stone-flies

eggs reach the bottom separately. From each egg hatches soon a tiny
flattened, soft-bodied, six-legged creature called a nymph, without wings
or wing-pads, and looking very much
like a Campodea (the simplest

living insect, see p. 61). This

crawls about, feeds, grows, moults,
nymph
grows, moults again and again (in a species observed by Lubbock there
were twenty-one moultings), and finally at the end of a year, or of two or
three years, depending on the species, is ready to issue as a winged adult.

During the nymphal life wings have been slowly developing, visible as
short pads projecting from the dorsal margins of the meso- and meta-thorax,
and appearing visibly larger after each moulting (Fig. 102). Respiration is

accomplished by flat, do not appear in some

leaf-like gills (Fig. 102) (these

species until after one or two moultings), arranged segmen tally along the
sides of the abdomen. The mouth-parts are well developed for biting
and chewing, with sharp-pointed jaws (mandibles). During its aquatic
life at the bottom of stream or pond the May-

fly has to undergo all the vicissitudes of an

exposed and protracted life; it is eagerly sought

after by larger, fierce, predaceous insects,

stronger of jaw and swifter than itself; it is

the prized food of many kinds of fishes, and it

has to struggle with its own kind for food and

place.
At the end of the immature life the nymphs
rise to the surface, and after floating there a
short time suddenly split open the cuticle along
the back and after hardly a second's pause

expand the delicate wings and fly away. Some

nymphs brought into the laboratory from a
watering-trough at Stanford University emerged
one after another from the aquarium with

amazing quickness. Almost all other insects

require some little time after the final moulting
for the gradual unfolding of the wings, and FlG I02
dry- . .
Young (nymph) of
ing and strengthening of the body- wall, before May-fly, showing (g) tracheal

flight or other locomotion. Most of the May-

fly species go through another moulting after ural size.)

acquiring wings, a phenomenon not known to occur in the case of any other
insect. The stage between the first issuance from the water with expanded
wings and the final moulting is called the subimago stage, and may last,
in various species, from but a few minutes to twenty-four hours. Such
is, in general, the life-history of the May-flies. As a matter of fact, the
life-history of no single May-fly species has yet been followed completely
68 The May-flies and Stone-flies

through. And here is an opportunity for some keen-eyed amateur ento-

mologist to add needed facts to our knowledge of insect life.
The breathing-organs of the nymph are of interest, as special adaptations
to enable them to take up oxygen and give off carbon dioxide without com-
ing to the surface, as do the water-beetles, water-bugs, mosquito-wrigglers,
and many other familiar aquatic insects. Each plate-like gill (Fig. 102)
is a flattened sac, with upper and lower membranous walls which run into

each other all around the free margin. Inside this sac is an air-tube
(tracheal trunk) with numer-
ous fine branches. By osmosis
an interchange of gases takes
place through the walls of the
tracheae and of the sac car-
bonic dioxide passing out, and
air from that held in solution
in the water passing in. If a

nymph held in a watch-glass

of water be watched, at times
all the gills will be seen rap-
idly vibrating, thus setting
up currents and bringing fresh
aerated water to bathe the
gills.
In the adult winged stage
(Fig. 103) the May-flies are
extremely frail and delicate-
bodied. The wings are fine
and gauzy, consisting of
the thinnest of membranes
stretched over a perfect net-
FIG. 103. May-fly, from California. (Natural size.) WO rk of veins. The fore
wings are always markedly larger than the hind wings; in some species
the latter are very small indeed, or even wanting altogether (Fig. 104).
The body- wall is weakly chitinized, and collected specimens almost always
shrivel and collapse badly in drying. The abdomen usually bears two
or three long filaments on its tip; the head is provided with compound eyes
and short awl-like antennae. The often-repeated statement in text-books
that adult May-flies have no mouth nor mouth-parts is not literally true
of all species, as weakly developed jaws and lips are present in some. But
they are in such weak and atrophied condition that they can hardly be func-
food. In
tional. probable, therefore, that no adult May-fly takes
It is

the males of some species the compound eyes present a very interesting
 The May-flies and Stone-flies

condition, being divided, each into two parts, by a narrow impressed line
or by a broader space The two
parts differ in the size of the
(Fig. 105).
facets of the ommatidia, i.e., eye-elements, and it has been ascertained (Zim-

merman, 1897) that this difference in size of facets

is accompanied by other and more important
structural differences, which make it certain that
the two parts of the eye have different powers of

seeing. One part is especially adapted for seeing

in the dark, or for detecting slight differences in

intensity of light, but is ill-fitted for exact sight,

while the part adapted for seeing in
other is

daylight, and for making a more exact picture of

outline. As the mating flights occur usually at
twilight or in the evening, Zimmerman believes
that this modification of the eyes of the males
is to enable them to discover the females in the

whirling shadow-dances. Chun has recorded a

similar division and difference in the eye of
certain ocean crustaceans and believes that the
"dark eyes" are used for seeing in the dimly FlG -
104. May-fly, Cmis
'

.. ,
. , .. ,
. ditmdtata, possessing only
lighted water below the surface, while the light one pair of wings. (Much
eyes" are for special use at the brilliantly lighted enlarged.)

surface. I have noted similar conditions in the eyes of both male and
female net-winged midges (Blepharoceridae), small, two- winged flies of

particularly interesting life (see p. 319). It is unusual to find such parallel

adaptations in forms so unrelated.
The May-flies show an anatomical condition of much interest to ento-

mologists in the paired openings for

the issuance of the eggs. Insects have
their organs arranged in pairs, one on
each side of the middle line of the
body, as the legs, wings, mouth-parts,
antennae, eyes, spiracles, etc., or exact-
ly on the middle line, as the heart,
FIG. 105. Section through head of alimentary canal, and ventral nerve-
male May-fly, Potamanthus brunneus,
cord. That is, the typical insect body
showing composition of compound
eye and two sizes of eye-elements is bilaterally symmetrical, and the
(ommatidia). (After Zimmer; greatly
more apparent this symmetry is the sim-
magnified.)
pler and more generalized the insect
is believed to be. All other insects but the May-flies have the two egg-

ducts, one from each egg-gland, fused inside the body, so as to form a short,
single, common duct on the median line. But the May-flies have the ducts
jo The May-flies and Stone-flies

separate; that is, paired and bilateral for their whole course. This is taken
to be an indication of the primitiveness and antiquity of the order.
If the May-flies are an ancient group of insects, and there is little doubt
of this, we have in them another example (we have previously noted one
in the case of Campodea, see p. 60) of primitive insects of excessively
frail and defenceless character persisting in the face of the strenuous struggle
for existence and of the competition, in this struggle, of highly developed,

specialized insect forms. Perhaps the solution of this problem in the case
of the May-flies is to be found in their extreme prolificness and in the

ephemeral character of their adult lives. It is only in the adult condition

that May- flies are so ill-fitted to defend themselves; so they simply make no

attempt to do so. They lay their eggs immediately on coming of age, and
thus accomplish the purpose of their adult stage. In their immature form
they are not so handicapped in the struggle for existence, although they
seem by no means in position to compete with some of their neighbors, like
the nymphs of the stone-fly and dragon-fly.
About 300 species of Ephemerida are known, of which 85 occur in
North America. Their classification has been comparatively little studied
and is a difficult matter for beginners. The differences among the adults
are so slight, and the preserved specimens are so uniformly misshapen
and dried up, that most of us will have to be satisfied with knowing that
we have in hand a May-fly, without being able to assign it to its genus.
Keys to the North American tribes and genera of May-flies may be found
by the student who may wish to attempt the generic determination of his
specimens, in a paper by Banks in the Transactions of the American Ento-
mological Society, v. 26, 1894, pp. 239-259.
There are better defined differences among the nymphs than among
the adults, but unfortunately the nymphs have been as yet too little studied
for the making out of a comprehensive key to the genera. Needham and
Betten give an analytical table of genera of Ephemerid nymphs as far as
known in the Eastern United States, in Bulletin 47 of the New York State
Museum, 1901.

ON THE under side of the same stones in the brook "riffles" where
the May-fly nymphs may be found, one can almost certainly find the very
similar nymphs (Fig. 106) of the stone-flies, an order of insects called

Plecoptera. More flattened and usually darker, or tiger-striped with black

and white, the stone-fly nymphs live side by side with the young May-flies.
But they are only to be certainly distinguished from them by careful exam-
ination. The gills of the immature stone-flies usually consist of single short
filaments or tufts of short filaments rising from the thoracic segments, one
tuft just behind each leg (Fig. 106), and not flat plates attached to the sides
 The May-flies and Stone-flies 71

of theabdomen as in the May-fly nymphs. The feet of the stone-flies have

two claws, while those of the young May-flies have but one. The stone-fly
nymph has a pair of large compound eyes, as well as three small simple eyes,
strong jaws for
biting and chewing (perhaps for

chewing Their nearest neighbors, the soft-bodied,

smaller May-fly nymphs!), and two slender back-
ward-projecting processes on the tip of the abdomen.
The legs are usually fringed with hairs, which makes
them good swimming as well as running organs.
The nymphs can run swiftly, and quickly conceal
themselves when disturbed.
nymphs, as far as known, require
All stone-fly
well aerated water; they cannot live in stagnant
pools or foul streams. Needham says that a large
number of the smaller species are wholly destitute
of gills absorbing the air directly through the skin.
;
FIG. 106. Young(nymph)
Nymphs brought in from a brook and placed in a
n-
vessel of still water will be seen with claws affixed,
-11 i stone-fly, from Can-

fornia. (Twice natural

vigorously swinging the body up and down, trying
siz e-)

to get a breath under the difficult conditions into which they have been

brought. The food-habits are not at all well known: some entomologists

assert that small May-fly nymphs and other soft-bodied aquatic creatures
are eaten, while others say that the food consists of decaying organic matter.
Here is another opportunity for some exact observation
by the interested amateur. On the other hand it is per-

fectly certain that the nymphs themselves serve as food

for fishes.
The fully worked-out life-history of no stone-fly seems
to have been recorded. The eggs, of which 5000 or 6000
may be deposited by a single female, are probably dropped
on the surface of the water, and sink to the bottom
after being,however, well distributed by the swift current.
Sometimes the eggs are carried about for a while by the
female, enclosed in a capsule attached to the abdomen.
The young moult several times in their growth, but
F F
'
*P r b a bly not nearly as many times as is common among
of nymph of stone- May-flies. When
ready for the final moulting, the nymph
fly. (Natural size.) cr awls out on a rock or on a tree-root or trunk on the

bank, and splitting its cuticle along the back, issues as a winged adult.
The cast exuviae (Fig. 107) are common objects along swift brooks.
The adults (Fig. 108) vary much in size and color, the smallest being
less than one-fifth of an inch long, while the largest reach a length of two
72 The May-flies and Stone-flies

inches. Some are pale green, some grayish, others brownish to black.
There are four rather large membranous, many-veined wings without pattern,
the hind wings being larger than the front ones. When at rest, the fore
wings lie flat on the back, covering the much-folded hind wings. The mouth-
parts are present and are fitted for biting, although the food-habits are not
known. It is asserted that some species take no food. The antennae are
long and slender. The abdomen usually bears a pair of long, many-seg-
mented, terminal filaments. The body is rather broad and flattened, and
there is no constriction between the thorax and abdomen. On the ventral

aspect of each thoracic segment there is a pair of small openings whose func-

FIG. 108. A stone-fly, Perla sp., common about brooks in California. (After Jenkins
and Kellogg; twice natural size.)

tion is unknown. The adults of certain species retain, although in shriveled

and probably functionless condition, the filamentous gills. This fact is of
importance in connection with the question as to whether insects are
descended from aquatic or terrestrial ancestors. Those* who believe in
I

the aquatic ancestry have found a simple origin for the spiracles (breathing-

pores) by imagining them to be the openings left when the gills, used in
aquatic life, were lost. But the adult stone-flies which retain their gills
also have wholly independent spiracles.
About 100 species of stone-flies are known in North, America. The
adults are to be found flying over or near streams, though sometimes
 The May-flies and Stone-flies 73

straying far away. They rest on trees and bushes along the banks. The
green ones usually keep to the green foliage, while the dark ones perch on

the trunk and branches. The various species are included in ten genera,
which may be determined by the following table:

TABLE OF NORTH AMERICAN GENERA OF PLECOPTERA.

The following technical terms not heretofore defined are used in this key: cerci,
slender processes projecting from the tip of the abdomen; radial sector, cubital vein,
and other names of veins in the wings may be understood by reference to Fig. 109.

P.A

FIG. 109. Diagram of venation of wing of a stone-fly; I, costal vein; 2, subcostal vein;
j, radial vein; 4, medial vein; 5, first anal vein; 6, radial sector; P, pterostigma;
A, arculus: a,, a 2 a z, apical cells. Between the medial and first anal vein is the
,

cubital vein, not numbered. CellM is the cell behind the medial vein; cell Sc is the
cell behind the subcostal vein.

A. With two long, many-jointed cerci.

B. Radial sector not reduced, i.e., with four or more branches.
C. Wings strengthened throughout by many cross-veins, there being many
cross-veins between the branches of the media, between the accessory
cubital veins, and in the anal areas of both pairs of wings. .PTERONARCYS.
CC. Wings with few or no cross-veins between the branches of the media,
between the branches of the cubital veins, and in the anal area.
D. Radial area of the fore wings with an irregular network of veins-
DlCTYOPTERYX.
DD.
Radial area of the fore wing with no cross-veins except the radial
cross-veins, or with a few regular cross-veins. .. .PERLA (in part).
BB. Radial sector reduced, i.e., with less than four branches.
C. Hind wings much broader than the fore wings.
D. With several cross-veins in cell M
of the fore wings.
E. Cell Sc of the fore wings with at least three cross-veins.
F. With three ocelli PERLA (in part).
FF. With only two ocelli PSEUDOPERLA.
EE.Cell Sc of the fore wings with only one or two cross-veins.
Small species of a green or yellow color CHLOROPERLA.
DD. With only one cross-vein in cell M
of the fore wings between the
arculus and the medio-cubital cross-vein CAPNIA.
CC. Hind wings of the same width as the fore wings; the anal area of the
hind wings not expanded ISOPTERYX.
AA. With the cerci rudimentary or wanting.
B. Second segment of the tarsi equal in length to the others; rudimentary cerci
present T^ENIOPTERYX.
74 The May-flies and Stone-flies

BB. Second segment of the tarsi small, shorter than the others, cerci absent.
C. Veins radiating from the ends of the radial cross-vein forming an X.
NEMOURA.
CC. Veins radiating from the ends of the radial cross-vein not forming an X.
LEUCTRA.

The genus Perla (Fig. 108) includes more species than any other. The
species of Pteronarcys retain gills in the adult condition. The species of
Chloroperla are small, delicate, and pale green. Leuctra includes the slender-
est of the stone-flies; they are small and brownish. Comstock says that
there are several species of stone-flies that appear on the snow on warm

days in late winter. They become more numerous in early spring, and
often find their way into houses. The most common one in Central New
York is Capnia pygmcea, which is grayish black. The
the small snow-fly,
female is 9 mm. (about f in.) long, with an expanse of wings of 16 mm.
(about -f in.), while the male is but 4$ mm. (about \ in.) long, and has
short wings which extend but two-thirds the length of the abdomen.
 CHAPTER VI
DRAGON-FLIES AND DAM-
SEL-FLIES (Order Odonata)

[EN it is on the mill-pond,

high noon
when leaves droop,and sun glares upon the
water, and the air is hot and still, when

other creatures seek the shade, and even

the swallows that skim the air morning
and evening are resting, then those other
swallows of the insect world, the dragon-
flies, are all abroad. One may stand
. . .

by the pond, and follow

side of a small
for hours with his eye the evolutions of one of the large dragon-flies skim-

ming over the surface in zigzag lines or sweeping curves, stopping still
and starting again, seeming never to rest, nor even to tire. Poised
in midair,

75
76 Dragon-flies and Damsel-flies

in the air, with the sunlight dancing on its trembling wings, it is indeed a
beautiful sight.

'"Dragon-flies? Folks call 'em devil's-darnin'-needles in our parts,

and they say they will sew up your ears.' Yes; and in some localities they
are called 'snake-doctors,' and are said
to bring dead snakes to life; and other

meaningless names are given them, such

as 'snake-feeders,' 'horse-stingers,' 'mule-
killers,' etc.; but in spite of all these

silly names and the superstitions

silly

they represent, dragon-flies are entirely

harmless to man are indeed to be
counted as friends, for they destroy vast
numbers of mosquitoes and gnats and
pestiferous little flies. To such creatures

they must seem real dragons of the air.

While one is standing by the pond let
him follow awhile the actions of a dragon-
that is making short dashes in different
fly
FIG. no. A dragon-fly (from life), directions close to the bank. Let him
fix his eye on a little fly hovering in the air, and note that after the

dragon-fly has made a dart toward it, it is gone. Let him repeat the
observation as the dragon-fly goes darting
hither and thither. It will be hard to see
the flies captured, so quickly it is done,
the place that once
'
but one can see that
knew them knows them no more.' And
the usefulness of the dragon-fly in taking
off such water-haunting pests will be
appreciated."
Thus entertainingly and truthfully writes
Professor Needham rf the strong- winged,

brilliantly colored, graceful insects of our

present chapter. If one could see through
muddy water and would fix his gaze on
the weed-choked slimy depths of the pond, FIG. in. The young (nymph) of
* dragon-fly. (From Jenkins and
he would see the dragon-flies in another
,,.,., ,.-. Kellogg; twice natural size.)
stage of their life, under very different
conditions of existence, and in very different guise. Crawling awkwardly
about over and through the decaying weeds and leaves and mud of the
bottom or lying in ambush, half concealed by coverings of slime,
would be seen certain strange big-headed, thick-bodied, dirty gray-green,
 Dragon-flies and Damsel-flies 77

wingless creatures from half an inch to two inches long. Occasionally

one of these creatures suddenly darts forward by spurting water from
the hinder tip of its body; occasionally one quickly thrusts out from
its head a vicious pincer-like organ which is more slowly withdrawn, or
rather folded up, with an unfortunate tiny water-animal squirming in the
toothed pincers. Still dragons, though now dragons of the deep instead of
flying dragons, these are our insects in their immature or larval life. Their

FlG. 112. Young (nymph) dragon-fly, showing lower lip folded and extended. (From
Jenkins and Kellogg; twice natural size.)

prey, consisting of water-bugs, May-fly larvae, small crustaceans, mol-

lusks, and any of the numerous aquatic insect larvae, including other
young dragon-flies, is probably always caught alive. Not by active
pursuit, as in the air above, but by lying in wait in the murky depths
of the pond until the unsuspecting insect comes within reach of the
extensible lower lip with broad spiny, jaw-like flaps at the
its pair of
clutching tip. The fierce young dragon, with its great
face of the
mouth and sharp jaws, is all concealed by this lip when folded up,
and there is little in the appearance of the dirty, sprawling, smooth-
faced creature to betray its dragon-like character. But appearances in
the insect world may be as deceptive as in our own, and too late the
careless water-bug out on a foraging swim for lesser prey finds himself in

range of a masked battery and becomes the preyer preyed upon.

About three hundred different species of dragon- and damsel-flies
(damsel-flies are the smaller, slender-bodied, narrow-winged kinds, see Fig.
113) are known in North America, about two thousand having been found
in all the world. In any single locality where conditions are at all favor-
able to dragon-fly life, that is, where there are live streams and ponds, from
a score to two or three times as many different dragon-flies can be found.
One hundred species occur in Ohio, and one hundred and twenty in New
York, states offering specially favorable natural conditions for them, while

only about fifty species have been found in California, a much larger but
more arid region. The young of no dragon-fly species is known to live in
salt water, although nymphs have been found in brackish water and in
 Dragon-flies and Damsel-flies

streams impregnated with sulphur from sulphur springs. Nor do dragon-

flies like cold weather. Although a few species are found in the far North
(recorded at 70 N. in Norway, 65 N. in Alaska, and 63 N. in Siberia)
and a few at high cold altitudes (as high as 10,000 feet) on mountain flanks,
the great majority of them need considerable temperature for growth and

development and even for activity during adult life. Calvert says that but
one species is known which regularly passes the winter in adult stage, and
that most dragon-flies live as adults from
but twenty-five to forty-five days, and
these in the summer. In California, where
the winter temperature at sea-level only
occasionally to 32 F., adult dragon-
falls

flies can be found in most of the months

of the year.
The adult dragon-flies are to be seen

pursuing their prey, like hawks, with

swift darting flights over ponds,
along
streams, and even scattered widely inland
over fields and in woods. A few kinds
have a liking for the vicinity of houses.
Needham, a careful student of these
has found that the hunting region
insects,
above and along the shores of a pond may
FIG. 113. Damsel-flies (narrow- be imaginarily divided into zones one
winged dragon -flies), (Natural above the other, each zone characterized
size; from life.)
by the presence of a few particular
dragon-fly species. "So, in fact," he writes, "we find the smaller damsel-
flies flying over the water in a straight course an inch or less above the

surface,and rarely venturing higher; the larger damsel-flies a little higher;

the amber wings at an average of about six inches; the larger skimmers
a foot or more '*rzzn the surface, and upland skimmers and darters still
higher. One has only to stand a little while by some small area of water
where all these are flying to see that each keeps rather closely to his proper
altitude. Why do damsel-flies keep so close to water? The reason is
not far to Dragon-flies eat one another the strong destroy the
seek.
weak. venture up into the altitude of the larger species means to run
If to

the risk of being eaten, we can readily see why the damsel-flies should

stay down below. The hawk may roam the air at will, but sparrows must
keep to the bushes."
We think of dragon-flies, as of albatrosses and Mother Carey's chickens,
as being always on the wing. They catch their prey while flying, eat it
while flying, mate while flying, and some of them deposit their eggs while
 Dragon-flies and Damsel-flies 79

on the wing. of course all dragon-flies rest sometimes, and some of

But
them, especially the damsel-flies, are at rest most of the time, clinging to
stems or leaves by the water's edge. The larger kinds may be found
occasionally perched on the tips of tall swaying reeds, or on a stump or
projecting dead limb. From these coigns of vantage they swoop like

a hawk on any rash midge that ventures awing in the neighborhood.

Cold or cloudy weather, or a strong wind, will drive most dragon-flies to
shelter.
The Odonata are unexcelled among insects for swiftness, straightness,
and quick angular changes in direction of flight. The successful main-
tenance of their predatory life depends upon this finely developed flight
function together with certain structural and functional body conditions
which might be said to be accessory or auxiliary to it. And this may be
an appropriate place to describe briefly a few of their salient structural
characteristics.
All dragon-flies have four well-developed wings, and all show such a
similar general bodily make-up and appearance, that from an acquaintance-

ship with two or three familiar species any member of the order can be
recognized as really belonging to the group. The body in all is long, smooth,
and subcylindrical or gently tapering. This clean, slender body offers
little resistance to the air in flight, and serves as an effective steering-oar.
The wings are long and comparatively narrow, fore and hind wings being
much alike, almost exactly alike indeed in the damsel-flies. The venation
is of the general type known as net-veining (Fig. 1146), the few strong longi-

tudinal veins being connected by many short cross-veins. The fore wings
are greatly strengthened along their costal (front) margin by having the
first longitudinal (subcostal) vein behind the 'margin placed at the bottom

of a groove, and the cross-veins in that groove so enlarged vertically as

to take on the character of flat, plate-like braces or buttresses. As, in
the figure-of-eight movement of the wing in flight, the front margin first
meets the resistance of the air, it is necessary that swiftly and strongly beat-
ing wings should be especially strengthened along this edge, and this is just
what the peculiar folding and bracing of the costal region of the dragon-fly's
fore wing accomplishes. .

The head is unusually large and is more than two-thirds composed of

the pair of great compound eyes. More than 30,000 facets have been
counted in the cornea of certain dragon-fly species, and this means that each
eye is made up of more than 30,000 distinct eye-elements or ommatidia,

each capable of seeing a small part or point of any object in range of vision.
Thus an image of a near-by object is made in fine mosaic, and the finer the
mosaic the more definite and precise is the vision by means of compound
eyes. These great eyes, too, have facets directed up and down and sidewise
80 Dragon-flies and Damsel-flies

as well as forward, and by a special sort of articulation of the head on the

thorax it can be rotated readily through 180, so that the principal part of
each eye can be directed sidewise or even straight down. For accurate
flight and successful pursuit of flying prey the dragon-fly has full need of

good eyes. It is to be noted, too, that the eyes are relatively largest in those

particular dragon-fly kinds which have the most powerful flight. On the
head, also, are three simple eyes (ocelli), the pair of very small awl-like
antennae, and the great mouth. The mouth is overhung as by a curtain
by the large flap-like upper lip (labrum). The jaws (mandibles) are strong
and toothed, and obviously well adapted for tearing and crushing the cap-
tured prey.
When the prey is come up with, however, it is caught not by the mouth
but by the "leg-basket." The thorax is so modified, and the insertion of
the legs such, that all the legs are brought close together and far forward,
so that they can be clasped together like six slender, spiny grasping arms

just
below the head. Although the catching and eating is all done in the
air and very quickly, observers have been able to see that the prey is caught
in this "leg-basket" and then held in the fore legs while being bitten and
devoured. These slender legs are used only very slightly for locomotion,
but they serve well for the light unstable perching which is characteristic
of the dragon-flies.
The internal anatomy is specially characterized, as might well be
imagined, by a finely developed system of thoracic muscles for the rapid
and powerful motion of the wings and the delicate and accurate move-
ments of the legs. The respiratory system is also unusually well developed,
such active insects needing a large quantity of oxygen, and generating a
large amount of carbon dioxide. The respiratory movements, according
to Calvert, consist in an alternate expansion (inspiration through the ten
pairs of breathing-holes, or spiracles, arranged segmentally on thorax and
abdomen) and contraction (expiration) of the abdomen. The rate of
movement varies greatly at different times owing to unknown causes, but
is always quickened by exercise, increased temperature, or mechanical irri-
tation. In different dragon-flies the inspirations have been noted to be
from 73 to 118 a minute.
The dragon-flies are famous for their beautiful metallic colors. As they
dart through the air one gets glimpses of iridescent blue and green and cop-

per, of tawny red and violet and purple reflections that are most fascinating
and tantalizing. Seen close at hand in the collections, however, they are
mostly dull-colored and, except for their "pictured" wings and the sym-
metry and trim outline of their body, rather unattractive "specimens." But
a freshly caught dragon-fly shows the real glory of the coloring: delicate
changing shades of green and violet and copper quiver in the great eyes;
 Dragon-flies and Damsel-flies 8 1

the thorax is translucent green or blue, and the long symmetrical

body is
warm red or deep blue or purple or green. It is often covered with a soft
whitish "bloom," that tones down the brilliant metallic iridescence. But
as the body dries, the colors fade. They are due not so much to pigment
as to the interference in reflection of the various color-rays, this interference

being caused by the structure of the body-wall. Just as soap-bubbles or

weathered plates of glass or mica produce brilliant colors by interference
effects, so does the semi-transparent laminate outer body-wall of the

dragon-fly produce its fleeting color glories. While the wings of many
kinds are clear, unmarked by blotches or line, the wings of others bear a
definite "picture" or pattern, usually light or dark brown or even blackish,
reddish, thin yellow, or whitish. These wing-patterns make the determination
of many of the dragon-fly species a very simple matter.
When the dragon-flies go winging about over ponds and streams they
are engaged in one of three things: in eating, in mating, or in egg-laying.
The prey of the dragon-fly may be almost any flying insect smaller than
itself, although midges, mosquitoes, and larger
flies constitute the majority

of the victims. Howard

says that the voracity of a dragon-fly may easily
be tested by capturing one, holding it by its wings folded together over its
back, and then feeding it on live house-flies. Beutenmuller found that
one of the large ones would eat forty house-flies inside of two hours. Howard
says that a dragon-fly will eat its own body when offered to it (query, to
its head ?) and that a collected dragon-fly, if insufficiently chloroformed and

pinned, will when it revives cease all efforts to escape if fed with house-flies,
the satisfying of its appetite making it apparently oblivious to the discom-
fort or possiblepain of a big pin through its thorax. That dragon-flies
are sometimes cannibalistic has been repeatedly confirmed by observation.
The nymphs have been seen to devour nymphs of their own and other

species; the nymphs of a European form have been observed to come out of
water at night and attack and devour newly transformed imagoes of the
same species, while several instances are recorded of the capture and devouring
of an imago of one species by an imago of another.
The good that is done by dragon-flies through their insatiable appetite
for mosquitoes is
very great. Now that we recognize in mosquitoes not
only irritating tormentors and destroyers of our peace of mind, but alarm-
ingly dangerous disseminators of serious diseases (malaria, yellow fever,
filariasis), any enemy of them must be called a friend of ours. A prize was
once offered for the best suggestions looking toward practicable means of
mosquitoes and house-
artificially utilizing dragon-flies for the destruction of
flies, but no very improvement on the dragon-fly's natural tastes
efficient

and practices were brought out by this essay competition.

In Honolulu, the principal city of our mid-Pacific territory, the mosqui-
82 Dragon-flies and Damsel-flies

toes are soabundant that no one neglects to enclose his bed carefully each
and all bedrooms are equipped with an ingenious
night in mosquito-netting,
canopy which can be folded closely in the daytime and readily spread over
the bed at night. The continuous and abundant presence of mosquitoes
is such a matter of fact that it has dictated certain particular habits of life
to the inhabitants of Honolulu. But in the daytime one is singularly free
from mosquito attack. Coincidentally with this one notes the surprising
abundance and strangely domestic habits of great dragon-flies. I have
watched dozens of dragon-flies hawking about a hotel lanai (porch) in the
heart of the town. No pond or stream is nearer than the city's outskirts.
Dragon-flies are in the main streets, in all the gardens, and they are chiefly
engaged in the laudable business of hunting the hordes of "day" mosquitoes
to their death. The most conspicuous features of insect life in Hawaii are
the hosts of dragon-flies by day and the hordes of mosquitoes by night. As
the dragon-flies unfortunately are not night flyers (although some forms

keep up the hunting until it is really dark), it is by night that one realizes
what a plague the mosquito is in the islands. Were it not for the dragon-
flies, life in the islands would be nearly intolerable. The rice-swamps and
taro-marshes and the heavily irrigated banana and sugar plantations offer
most favorable breeding-grounds for the mosquitoes, but also fortunately
for the dragon-flies as well. The mosquitoes of Hawaii are not indigenous;
they were introduced with white civilization. It is told, and is not improb-

able, that the skipper of a trading schooner in early days, to revenge himself
for some slight put on him by the natives, purposely put ashore a cask of
water swarming with mosquito wrigglers. It needed no more than that
to colonize this fascinating tropic land with the mosquito plague. How
the saving dragon-fliescame is not yet come to be tradition; indeed, few
Hawaiians understand how important a part the dragon-fly plays in their
life. They do appreciate the mosquito.
In the Samoan Islands, too, where we have another tropical colony,
the mosquitoes are a great plague. Here the matter is made more serious.
The Samoan mosquitoes are carriers and disseminators of a dreadful disease
known as elephantiasis from the enormous enlargement of the legs and
arms of sufferers from it. This disease is the great scourge of these islands,
more than 30% (from my own observation; 40% and 50% are estimates
given by other observers) of the natives having it. (For an account of
the role of mosquitoes in the dissemination of malaria, yellow fever, and

elephantiasis, see Chapter XVIII of this book.) The dragon-flies are, in

Samoa as in Hawaii, conspicuous by their abundance and variety, and they
do much to keep check the quickly breeding mosquitoes.
in

Watching the dragon-flies over a pond, you may occasionally

flying
see one poising just over the surface of the water, and striking it with the
 Dragon-flies and Damsel-flies 83

tip of the abdomen; or another kind may be seen to swoop swiftly down to
the surface occasionally in its back-and-forth flight, and to dip the tip of

FIG. 1146.
Stages in the development of the giant dragon-fly, Anax junius. a, youngest stage; b,
c, and d, older stages, showing gradual development of the wings. (Young stage,
slightly enlarged after Needham; adult three-fourths natural size.)

the body moment into the water. These are females engaged in laying
for a
their eggs. The eggs issue in small masses, usually held together by a gelat-
inous substance. From several hundred to several thousand eggs are laid by
 Dragon-flies and Damsel-flies

each female. Needham counted 110,000 eggs in a single egg-mass of Libellula.

Sometimes the eggs may be laidon wet mud or attached to moist water- or
shore-plants. The damsel-flies and a few of the dragon-flies insert the eggs
in the stems of dead or living water-plants below the surface of the water.
To do this they have to cling to the stem, with the abdomen or sometimes
the whole body under water, and cut slits in it with the sharp ovipositor.
The eggs are sometimes laid on submerged timbers and moss- or alga-covered
stones. Kellicott observed females of A rgia putrida (a damsel-fly abundant

along Lake Erie) to remain wholly under water for from five to fifty-five
minutes at a time. These females were accompanied by males which also
stayed under for similar lengths of time.
The eggs hatch after various periods, depending on the species of dragon-
fly and on the time of year of oviposition. In midsummer Needham found
the eggs of some species to hatch in from six to
ten days, while others laid in autumn did not hatch
until the following spring. In the same
lot of eggs

the period of incubation

may vary even in midsum-
mer from a week to more than a month.
From the eggs come tiny, spider-like nymphs
with long, slender legs, thin body, and no sign of
wings. Even in the largest dragon-fly species the
just-hatched young are only about one-twelfth of
an inch long, while the nymphs of the common
Libellulas are only one-twenty-fifth of an inch long
at hatching. They begin their predatory life, con-

fining their attention at first to the smaller aquatic

creatures, but with increasing size and strength
and confidence being ready to attack almost any of
the under-water dwellers. Even fish are seized by
FIG. The
115. young
of a damsel-
the larger nymphs, Needham having seen the
(nymph)
(narrow-winged dra- nymphs of one species seize and devour young
fly
gon-fly), Lestes sp. The brook-trout as long as themselves.
three leaf -like processes
at the tip of the abdo- The young of different species differ consider-
men are gills (Twice ably in size, shape, and duration of their nymphal
natural size.)
existence. The nymphssome species require
of

more than a year some others are ready

to develop into adults, while those of
to transform in a few months, not a few dragon-fly species having two gener-
ations a year. The one-year life cycle, however, is usual among the more
familiar dragon-flies, the eggs laid during midsummer hatching in late sum-

mer, the nymphs hibernating and being ready to emerge the following sum-
mer. Needham thinks that the damsel-flies have a number of broods in
a season, the processes of transformation and oviposition beginning as soon
 Dragon-flies and Damsel-flies 85

as the weather permits, and continuing industriously to the close of the

season.
The nymphs cast the skin repeatedly during their growth and develop-
ment, although the exact number of moultings is not known for any species.
After two or three moults the wing-pads appear and with each successive
moult increase in size. Immediately after moulting the nymphs are light
greenish or gray, and their characteristic color pattern is distinct, but they

gradually darken, the pattern becoming more and more obscure until by
the t'me for another moulting the body is uniformly dark and dingy. The

nymphs (Fig. 115) of the damsel-flies are elongate and slender, and have
three long conspicuous gill-plates at the tip of the abdomen, which they
can also use as sculls for swimming. The dragon-fly nymphs are robust-
bodied, some of them indeed having the abdomen nearly as wide as long
and much flattened. All the nymphs are provided with the long grasping
lower which can be folded mask-like over the face when not engaged
lip,
in seizing prey. The mandibles are strong and sharp and the whole mouth
is well fitted for its deplorable but necessary business.
The true dragon-fly nymphs do not have plate-like gills, like those of the
damsel-flies, nor any other external kind, but have the posterior third of
the intestine lined with so-called internal gills. These internal or rectal
gills are in six longitudinal bands, each consisting of two thin rows of small
plates or tufts of short slender papillae. Water is taken into the intestine
through its posterior opening and, after bathing the gills, giving up its dis-

solved oxygen, and taking up carbon dioxide, it is ejected through the same

opening. When this water is ejected violently it serves to propel the nymph
forward. It is also apparently occasionally used for defence.

Just as the adult flying dragon-flies keep to certain regions above or

in the neighborhood of the pond, so Needham has found the nymphs to
have various preferred lurking-places in the pond. The damsel- fly nymphs
and a few of the more active dragon-fly nymphs clamber among submerged

vegetation or inhabit driftwood and submerged roots or brush. The heavier

sprawling Libellulid nymphs usually crawl over the bottom or climb over
fallen rubbish, while certain other Libellulids and some similar forms occupy
the mud or sand of the bottom. The nymphs of one of these latter kinds
isdescribed as each scratching a hole for itself and descending into it like
a chicken into a dust-bath, kicking the sand over its back and burrowing
until all but hidden, only the tops of its eyes, the tips of its treacherous labium,
and the respiratory aperture at the end of the abdomen reaching the surface.
After the few weeks or month or year which the nymph requires for its full
growth and development it is ready to transform. If in early summer, when
the dragon-flies are beginning to appear, one will go out to the dragon-fly

pond a little after daylight, he will see this transforming or issuance of the
86 Dragon-flies and Damsel-flies

winged imagoes busily going on. The nymphs crawl out of the water, and
up on stones or projecting sticks, or on bridge-piles or the sides of boats,
or on the stems of weeds growing by the water's edge. Here they cling quietly,

awaiting the moment when the chi-

tinous body-wall shall split lengthwise

along the back of the thorax, and the

made-over body inside with its damp,
compressed wings, its delicate trans-

parent skin, and changed mouth-parts

and legs shall slowly work its way out
of the old nymphal coat. The nymphs
of some and damsel-flies
dragon-flies
FlG. 116. The issuance of an adult white crawl out among theweeds and grass
tail, Plathemis trimaculata, (After Need- of the shore for some distance before
ham; natural size.)
choosing a resting-place, and none of
these will be very readily seen. But careful searching in a place from which
winged individuals are occasionally arising will soon reveal the transforming
in all of its stages (Fig. 116). It takes some time for the emergence of the

damp, soft imago from the nymphal skin, and some further time for the
slow expanding and drying of the wings, and the hardening of the body-
wall so that the muscles can safely pull against it. When all this has come
about the imago can fly away. But even yet the colors are not fully acquired

FIG. 117. Adult and last exuvia of the whitetail, Plathemis trimaculata.
(Natural size.)

and fixed, and these fresh imagoes have an unmistakably new and shiny
appearance. They are called teneral specimens. Usually the emergence
of nymphs from the pond and the subsequent transforming cease by the
middle of the forenoon, and after that one can find only the frail, drying
 Dragon-flies and Damsel-flies

cast nymphal skins or exuviae, clinging here and there to stones and plant-
stems. Attached to these exuviae there may be often noted two or three short,
white, thread-like processes. These
are the dry chitinous inner linings
of the main tracheal trunks of the

dragon-fly which were moulted with

the outer body- wall. As the main
tracheal tubes are really imagina-
tions of the outer skin, it is obvious
that the inner lining of the trachea
is continuous with the outer coat
(chitinized cuticle) of the body-wall
and so is naturally cast off with it.

Although the habits of the adult

dragon-flies must be studied out of
doors, the nymphs can be brought
indoors and kept alive so that their
Adult and exuvia of the damsel-
walking and swimming and hiding FIG. 118. last
, . Lestes uncata. (Natural size.)
and capturing of( prey, and often
, ,, fly,

their transformation into winged imagoes, can be readily observed. In

their natural habitat some of these observations are nearly impossible,
and for school-room or private-study aquaria

hardly any other animals can be found of

more interest to the observer, whether child or
grown-up, than the dragon-fly nymphs.
Professor Needham, who has done more
and better work in the study of the immature
life of dragon-flies than anybody else, gives
the following directions for collecting and
rearing the nymphs:
"If one wishes to collect the nymphs which
liesprawling amid fallen trash, a garden-rake
with which to draw the trash aside, fingers not
too dainty to pick them up when they make
themselves conspicuous by their active efforts
to get back into the water, and a pail of
water in which to carry them home, are all

FIG. 119. A home-made water- the apparatus required,

net for collecting dragon-fly "A rake will bring ashore those other
nymphs. (After N
nymphs whkh bum)W
shallowly under the
sediment that lies on the bottom, and also a few of those that cling to vegeta-
tion near the surface; but for getting these latter a net is better. Fig. 119
88 Dragon-flies and Damsel-flies

shows the construction of a good water-net that can be made at home out
of a piece of grass-cloth, two sizes of wire, and a stick.
"The best places to search for dragon-fly nymphs in general are the
reedy borders of ponds and the places where trash falls in the eddies of
creeks. The smaller the body of water, if permanent, the more likely it
is to yield good collecting. The nymphs may be kept in any reasonably
clean vessel that will hold water. Some clean sand should be placed in
the bottom, especially for burrowers, and water-plants for damsel-fly nymphs
to rest on. They may be fed occasionally upon such small insects (smaller
than themselves) as a water-net or a sieve will catch in any pond. Their
habits can be studied at leisure in a dish of water on one's desk or table.
"The best season for collecting them is spring and early summer. April
and May are the best months of the year, because at this time most nymphs
are nearly grown, and, if taken then,
will need to be kept but a short time
before transforming into adults. And
this transformation every one should

see; it will be worth a week's work at

the desk; and as it can be appreciated
only by being seen, some simple direc-
tions are here given for bringing the
FIG. 120. A simple aquarium for rear- to ma turity. Place them in a
Need- nymphs
*
ing dragon-fly nymphs. (After
ham.) wooden pail or tub (Fig. 120). If

the sides are so smooth that they cannot crawl up to transform, put some
sticks in the water for them to crawl out on. Tie mosquito-netting tightly over
the top, or, better, make a screen cover; leave three or four inches of air
between the water and the netting; feed at least once a week, set them where
the sun will reach them; and after the advent of warm spring weather look
in on them early every morning to see what is going on."
Elsewhere Professor Needham says that nymphs may be fed bits of
fresh meat in lieu of live insects. If meat
must be kept in motion
is fed, it

before them, as they will refuse anything that does not seem alive. Some
nymphs will take earthworms. Care must be taken to keep cannibalistic
kinds apart from others. When the nymphs transform the freshly issued
imagoes should be transferred each with its cast skin (exuvia) to dry boxes
for a short time, till their body-wall and wings gain firmness and the colors
are matured. The imago and its exuvia should always be kept together.
Specimens of the adults for the cabinet should have the wings spread
and moths (for directions for spreading see the Appendix).
like butterflies
The slender and brittle dried abdomen breaks off very easily, and a bristle
or fine non-corrosive wire should therefore be passed lengthwise through
the body as far as the tip of the abdomen. A
couple of insect-pins, inserted
 Dragon-flies and Damsel-flies 89

lengthwise one at each end of the body, are used by some. Specimens
intended for exchange should not be pinned up, but "papered," i.e., put
with folded wings into an enclosing little triangular paper envelope made

by folding an oblong paper sheet once diagonally and then folding over
slightly the
two margins.

/.!,
.,
''

10 9
FIG. 121. Diagram of venation of
wing of dragon-fly, o, antecubitals; b, postcubitals;
AT, nodus; P, pterostigma; A, arculus; /, triangle. (After Banks.)

TABLES FOR CLASSIFICATION.

KEY TO SUBORDERS (IMAGOES).

Front and hind wings nearly similar in outline, and held vertically over the back
when at rest; head wide and with eyes projecting and constricted at base.
(Damsel-flies.) Suborder ZYGOPTERA.
Front and hind wings dissimilar, hind wings usually being much wider at base, and
both pairs held horizontally outstretched when at rest; eyes not projecting
and constricted at base (Dragon-flies.) Suborder ANISOPTERA.
KEY TO SUBORDERS (NYMPHS).
Posterior tip of abdomen bearing three, usually long, leaf-like tracheal gills.

(Damsel-flies.) Suborder ZYGOPTERA.

Posterior tip of abdomen with five, converging, short, spine-likeappendages.
(Dragon-flies.) Suborder ANISOPTERA.

SUBORDER ZYGOPTERA.
KEY TO FAMILIES (IMAGOES).

Wings with not less than five antecubital cross-veins (Fig. 121).
Family CALOPTERYGID.E.
Wings with not more than three, usually two, antecubitals (Fig. 121).
Family AGRIONID<E.
KEY TO FAMILIES (NYMPHS).
Basal segment of the antennas extremely elongate Family CALOPTERYGID-E.
Basal segment of the antennae short, subrotund Family AGRIONID^E.

The family Calopterygidae includes but two genera, Calopteryx, in which

the basilar space of the wings is open and the wings themselves are rather
broad near the tip, and Hetaerina, in which the basilar space is net-veined
and the wings narrow.
Calopteryx maculata (Fig. 122), the most familiar representative in the
Eastern States of the first genus, has velvety black spoon-shaped wings,
9 Dragon-flies and Damsel-flies

(brownish in freshly moulted, or teneral specimens), and a long, slender body,

of striking metallic blue or green. The females can be distinguished from
the males by their possession of a milk-white pterostigma These
(Fig. 121).
beautiful "black wings" are found 'n gentle fluttering flight,
usually along
small streams in woods or meadows. The female lays her eggs "among
the rubbish and mud along the
borders of ditches," and the
nymphs found in the ditches
and streamlets have the middle
one of the three caudal gills flat
and shorter than the other two.
Kellicott has seen the males of
this species fight fiercely with
each other. " Two will fly about
each other, evidently with con-
suming rage, when one finally
appears to have secured a posi-
tion of advantage and darts at
his enemy, attempting, often suc-
FIG. 122. The black wing, Calopteryx cessfully, to tear and damage
maculata.
his wings."
The best known representative of the other genus is a perfect master-
piece of insect beauty and grace. Entomologists know it as Hetcerina

americana (Fig. 123); I suggest that we call it the "ruby-spot," although

only the males bear the gem. The head and thorax of the males are
coppery red, the abdomen me-
tallic green to coppery, and the
basal fourth of each of the long,
slender, and otherwise clear wings
isbright blood-red. In the females
the whole body is metallic green,
with the basal third of the wings
pale yellowish brown. These dam-
sel-fly beauties are shy
and retiring,
rarely venturing more than a few
feet away from the willow-overhung
bank of their favorite swift-running
stream. Sometimes hundreds of FIG. 123. The ruby-spot, Hetcerina
them come together and cling in americana.

graceful festoons to the drooping willow branches. Then they look like
strings of rubies, or of warm red flowers or seeds.
The family Agrionidae includes the host of slender-bodied, narrow- and
 Dragon-flies and Damsel-flies 91

clear-winged true damsel-flies. Most of them are small, and many keep
so closely in low herbage or shrubby woodland that they attract little atten-
tion. A few of the longer-bodied and longer-winged forms, however, fly
in the open along the stream-banks or over the ponds. Some are strikingly
varied with black and orange or yellow, and all, whether brightly colored
or dull, are graceful and charming. There are at least a dozen genera of
Agrionids in this country, comprising about seventy-five species, but their
classification is too difficult to be undertaken by general students. Damsel-
fliesdeposit their eggs in the tissue of aquatic plants by cutting slits in the
stems with their sharp ovipositor. The nymphs are slender and elongate,
and can readily be known by the three caudal leaf-like tracheal gills. The
nymph stage of these forms is much shorter than with the true dragon-flies,

lasting usually probably but a few weeks, or at most two or three months.
When ready to transform the nymphs crawl out of the water and into the
low herbage on the stream or pond bank. I have seen scores of freshly

emerged damsel-flies rising from a few square yards of tall grass near a pond,
although it required close search to discover the nymphs, so well concealed
were they in the dense tangle.

SUBORDER ANISOPTERA.
KEY TO FAMILIES (IMAGOES).

Antecubitals of the first and second rows mostly meeting each other; triangle of
fore wings with long axis at right angles to the length of the wings, triangle
of hind wing with long axis in direction of the length of the wing.
LlBELLULID^E.
Antecubitals of the first and second rows not meeting (or running into each other)
except the first and another thick one; triangles of fore and hind wings of
similar shape (Fig. 121).
Eyes meeting above on middle line of head; abdomen with lateral ridges.

Eyes just touching at a single point or barely apart; abdomen without lateral

ridges ...................................... CORDULEGASTERID^:.

Eyes distinctly separated; abdomen without lateral ridges ...... GOMPHID^E.
KEY TO FAMILIES (NYMPHS).
Under-lip (labium) flat, not concealing most of the face, with jaw-like or oblong
side pieces (lateral lobes).
Antennae 7 -segmented, tarsi 3-segmented, climbing nymphs. .^ESCHNID^:.
Antennas 4-segmented, the 'fourth segment rudimentary; fore tarsi 2-seg-
mented; burrowing nymphs .......................... GOMPHID.E.
Under-lip (labium) spoon-shaped, covering most of the face, when closed, with nearly
triangular side pieces (lateral lobes).
Two stout teeth with a notch between them on the middle lobe of the under-

lip(labium) ................................. CORDULEGASTERIDJ;.

A single median tooth on the middle lobe of the under-lip ---- LIBELLULID^E.
92 Dragon-flies and Damsel-flies

The family Cordulegasteridae includes only seven species of dragon-flies

found in the United States, all belonging to one genus, Cordulegaster. They
are large, with eyes barely touching on top of the head, without metallic
body-colors, and with clear wings. The nymphs burrow into the sand or
vegetable silt on the bottom of shallow places. Thus buried, with only
the top of the eyes and tip of the abdomen showing, they remain motionless
for a long time, if prey does not come near. "In a dish of sand on my table,"
says Needham, "I have had a nymph remain without change of position
for weeks,no food being offered it. Let any little insect walk or swim near
the nymph's head, and a hidden labium springs from the sand with a mighty

sweep and clutches it." The imagoes are strong flyers and have the habit
of flyingback and forth, as on a regular beat, over some small, clear stream.
Thefamily Gomphidas includes six genera, comprising about fifty species
in our country. They are mostly large forms, clear- winged and with bodies
striped with black and green or yellow. They are readily distinguished
by the wide separation of the rather small eyes. The abdomen is stiff and
spike-like. The
eggs, held in a scanty envelope of gelatin, are deposited

by the repeated descent of the flying female to the water of a clear pond
or flowing stream, the tip of the abdomen first striking the surface. The
gelatin dissolves and the
eggs, scattering, sink to the bottom and become
hidden in the silt. The nymphs
are active burrowers, capturing their prey
either on or beneath the surface of the bottom silt. The adults often alight
on foliage, or on the surface of some log stretching across a stream, or on
the bare soil of a path or roadway. They do not fly about in apparent
sportiveness as the skimmers (Libellulidae, p. 95) do, nor, like the skim-
mers, perch atop a slender twig. June is the best month in the East for
these dragon-flies. The principal genus of the family isGomphus, which
includes one-third of all our Gomphidae. Of these Gomphus exilis is
probably the most common one in the Northeastern States. Its head is pale
green, thorax brownish with two oblique green bands on each side, and
abdomen blackish brown with a basal green spot or band on the back
.

of each segment. The nymphs transform at the very edge of the water
?

seldom crawling more than an inch or two above it.

Hagenius brevistylus
is a large black-and-yellow species common in the East, South, and Middle
West. The nymph has an unusually wide, flattened body.
The ^Eschnidae include our largest, swiftest, and most voracious dragon-
flies. Various species are flying through the whole season from early spring
to late summer. Some roam far from water, being found over dry fields
and roadways, and even in houses. Some forms fly until late in the even-
ing, making life a burden
for the mosquitoes gathering for their
night's
singing and feasting. The eggs are thrust into the stems of aquatic plants,
in floating timbers, in the wood of piers, etc., at or near the surface of the
 Dragon-flies and Damsel-flies 93

water. The nymphs are slender, clean creatures, with smooth bodies pat-
terned with green and brown, and very active, strong, and brave. They
climb among green plants and roots or submerged driftwood along the border
of open water or the edge of a current. The imagoes of this family can be
recognized by the meeting of the eyes all along the top of the head. The
wings are long, broad, and clear, and the body-colors are mostly bright blue
and green. The family is represented in the United States by about twenty-five
species, belonging to six genera. Anax junius, one of the commonest dragon-
over the United States, and found also from Alaska to Costa Rica,
flies all

in China, Siberia, and in various islands of the Pacific, notably the Hawaiian

group, is the most inveterate enemy that the mosquito has. It is conspicu-

ously on the wing from early spring to

flying from daylight to dark,
late fall,
and doing untold good by its ceaseless
warfare on the mosquito hosts. It
can be recognized by its clear wings,
large size (wings over two inches long),
, , . , , j
,, , , -i FIG. 1240. FIG. 1246.
and bright-green thorax and head, the
latter bearing on thee nnner front
FlG ' I2 4 a - T P of head showing charac-
>

upper I a 1 -

teristic mark n front Q{ eyeg) of Anax

round black spot surrounded by yellow, junius. (Enlarged.)
Fl of head showing charac-
the yellow
'
encircled by a dark-blue f- p^.-Top
teristic mark in front of eyes, of ALschna
ring (Fig. 1240). A still larger member constricta. (Enlarged.)
of this family is the great "hero"
dragon-fly, Epiaschna heros, like Anax junius in general appear-
which is
ance, but has wings two and one-half inches long, and abdomen nearly three
inches long. It has a black T
spot on the upper face, instead of a round
one. Another similar, widely distributed and common form is ^Eschna
constricta, about the size of Anax junius, reddish brown marked with bright
green, and with a black T spot on the upper front of face (Fig. 1246). The
males have the abdomen marked with blue, with little or no green, while
the females have but little blue or none at all.
The members of the family Libellulidae are called "skimmers." They
may be seen continually hovering over the surface of still water, or swiftly
foraging over fields. Many of them have the wings strongly marked with
large black or brown or milk-white blotches, and the abdomen is often
covered with a whitish powder or "bloom." They outnumber all the other
true dragon-flies in point of species, and except for Anax junius, ^Eschna

constricta, and perhaps the giant hero dragon-fly, include the most familiar
and wide-spread members of the order. One of the best known and most
beautiful of the skimmers is the pond-loving "ten-spot," Libellula pulchella

(Fig. 125), found all over the country. Each of its wings has a longitudinal
basal blotch, a median blotch (at the nodus), and an apical blotch of black-
94 Dragon-flies and Damsel-flies

ish brown. The males have the space between these blotches milky white.
In old individuals the abdomen has a strong whitish bloom. Other familiar

FIG. 125. The ten-spot dragon-fly, Libellula pulchella. (After Needham; nat. size.)

and well-marked species of Libellula are L. basalis, with blackish-brown body

and with the basal third to half of the wings dark brown or black and the
rest of the wing clear, or in the old males chalky white out as far as the

FIG. 126. Libellula semi-fasciata. (After Needham; natural size.)

pterostigma, and in the females with brownish apices; L. quadrimaculata,

with olive or yellowish body marked with black, front wings with more
 Dragon-flies and Damsel-flies 95
or less yellowish at baseand along the front margin, and a small fuscous
nodal spot, hind wings with a yellowish-black triangular basal spot and
fuscous nodal spot; and L. semi-fasciata, whose complex wing-markings are

FIG. 127. The water-prince, Epicordulia princeps, female.

(After Needham; natural size.)

shown in Fig. 126. Tramea is a genus of large swift dragon-flies whose

hind wings have the base expanded and conspicuously colored. Tramea
lace-ata is a familiar species. The water-prince, Epicordulia princeps (Fig.

FIG. 128. The amber wing, Perithemis domilia, male at left, female at right.
(After Needham; natural size.)

127), is a common large dragon-fly, but one hard to capture because of its
fine flight. The wings show a basal patch, often nearly wanting on the
front pair, a patch at the nodus, and a black apex. It likes "ponds or slug-
96 Dragon-flies and Damsel-flies

gish streams with muddy reed-grown banks, and seems absolutely tireless
in flight; very rarely indeed is one seen resting." One of the smallest of

FIG. 129. The wind sprite. Celithemis eponina. (After Needham; natural size.)

FIG. 130. Tetragoneuria epinosa, female. (After Needham; natural size.)

the true dragon-flies is the amber wing, Perithemis domitia (Fig. 128). The
wings are clear amber, unmarked in the male, but richly spotted with dark
 Dragon-flies and Damsel-flies 97
brown in the female. It has a slow hovering flight and often rests on the
tips of erect reeds with wings held perfectly horizontal. It is only on wing
in quiet, warm sunshine; clouds or cold breezes send them quickly into
hiding. Among the familiar Libellulids with unblotched wings is Meso-
themis simplicicollis, an abundant species east of the Rockies. The
females and young males have head, thorax, and front half of abdomen
green, the hinder half blackish brown. In old males the body becomes
grayish blue with a whitish bloom. Williamson says that sometimes two
males will flutter motionless, one a few inches in front of the other, when
suddenly the rear one will rise and pass over the other, which at the same
time moves in a curve downwards, backwards, and then upwards, so that
the former position of the two is just reversed. These motions kept up

FIG. 131. The whitetail, Plathemis lydia. (After Needham; natural size.)

with rapidity and regularity give the observer the impression of two inter-
secting circles which roll along near the surface of the water.
The whitetail, Plathemis lydia (Fig. 131), resembles the ten-spot, but
is one-fourth smaller. In the males also the apex of the wings is usually
clear, not brown. The whitetail rather likes slow-flowing brooks and
open ditches. When alight it has the habit of setting its wings aslant down-
ward and forward with a succession of jerks. Needham thinks that the
powdery whiteness of the body of the old males (in females and young males
the body is brown marked with yellow) must render it more easily seen by
its enemies, the king-birds and others, and thus be a
disadvantage in the
struggle for existence. He says, indeed, that the whitest ones avoid rest-
98 Dragon-flies and Damsel-flies

ing-places over a dark background and settle oftenest on white sticks, on

bleached stumps, or on light-colored earth. Very frequently one will alight
on a white insect-net when it is laid down, or even when still held in the
hand.

IM. H.W.I
 CHAPTER VII

THE TERMITES, OR WHITE ANTS (Order Isoptera)

NCE when camping in the King's River

Canon, one of the great vertical -walled, flat-
floored canons of the Sierra Nevada, the
boldest axeman of our party attacked the
fallen trunk of a once towering yellow pine.
The practical outcome of this attack was
a sufficient supply of firewood for the
cook's stone-built stove, but the great log

yielded better things than chips and chunks.

A few blows showed it to be the home. of
a thrivingcolony of the largest of the
American termites (Termopsis angusticollis), and the thousands of indi-
viduals in this insect household were objects of interested observation
the summer through. We had heard of the rarity of white-ant queens in
collections,and saw in this isolated and apparently easily "rounded-up"
community an easy chance to discover the egg-laying queen of this species.
But we had not reckoned with the Californ'a manner of tree-trunk: it
outlasted the summer's chopping by two score feet of log four feet thick.
Yellow pines grow 250 feet high in the Sierran forests. But although
no queen was found, the make-up of the buried termite city was revealed.
Galleries and chambers, secret ways and narrow tunnels were all ex-

posed, and the interesting communal life of these soft, white-bodied little
creatures was made partly known to us.
We have in the United States but few kinds of termites, and these
much less interesting in habit than those of tropic lands. The Amazons
and Central Africa are the centers of termite life, and there, because of their

great mounds, their serious ravages on allthings wooden, and their enor-
mous numbers, the white ants come to be nearly the most conspicuous of
the insect class. Drummond's account, in his Tropical Africa, of the habits
and of the termites of the Central African region is simply a tale of
life

marvels. And the scattered accounts of the Brazilian species are hardly
less wonderful. In the South Sea, too, the termites play their part promi-
99
IOO The Termites, or White Ants

nently. I have seen scores of cocoanut-palms in Samoa with their trunks

traced over from ground to "feather-duster" top, a hundred feet above, by
the laboriously builded wood-pulp tunnels of the termites. Each of these
trees carried also on its trunk, about four feet from the ground, a termite
"shed" or depot (Fig. 133), a foot thick, a foot wide, and two feet long,
made, like the tunnels, of pellets of chewed wood, glued together with saliva,
and filled with crowded galleries and chambers.

FIG. 132. Giant hillock-nests of termites in tropical Africa.

(Adapted from Drummond.)
But in the United States our few species make their communal nests
in dead and dying wood, or underground, and not being given to building
great dome-like mound-nests, or making covered ways up all the trees of
a great forest or plantation, are not as conspicuous as their tropical cousins.
Still, few observers of insects have failed to notice the little, white, wingless
worker termites, scurrying about when some dead stump is overturned or
splitopen, or to see the winged males and females swarming out of the
ground some sunny day, and, after a brief period of flight, pursued by birds
and predaceous insects, settling to earth again and losing their wings.
Before proceeding to take up the incompletely known life-history of our
American termites it will be advisable to describe their
general structural
 The Termites, or White Ants 101

characters and the composition of the termite communities. The body

is always soft, and usually milky-whitish in color, though sometimes light
or dark brown. It is plump, and slightly broader than thick. The abdo-
men is joined broadly to the thorax, not by a little stem or peduncle as in
the ants, with which insects the name "white
ants" long and widely
(unfortunately too
used to be done away with) confuses the
ermites in the popular mind. The termites
not only are not ants, but are neither nearly
related to them nor of similar structure.
The only resemblances between the two forms
exist in the communal life and in the com-
position of the community by different kinds
of individuals. The termites are either blind
or have only simple eyes, have slender an-
tennas which look as if made up of tiny beads

strung a-row, and have biting mouth-parts

with strong jaws. They live in small or large
communities, the individuals in any one of
which, although belonging to the same species,
being of from three to eight different kinds
or castes. That is, each community is com-
posed of winged and wingless individuals,
the winged being males and females, while
the wingless include immature individuals,
sexually incomplete workers and soldiers,
and also so-called complemental males and
females which are individuals able to help
in the increase of the community. In some
species there are no workers, while in others
the workers may be of two kinds. The
FIG. 133. Termite shed on
soldiers differ from all the others in he
cocoanut-palm in Samoa. From
extraordinary development of their the shed note numerous tunnels
jaws,
leading down to the ground, in
which are long and scissor-like; their heads which is the main nest of the
are also much enlarged and strongly chitin- community; a few tunnels (only
one visible in the picture) lead
ized. The food of all consists mainly of
up the trunk of the tree. (Pho-
dead wood, and of curious pellets excreted tograph by the author.)
from the intestine and called "proctodeal
food." In addition some species attack live .wood and even soft plants,
and cloth, books, papers, etc., suffer from termite ravages. The serious
nature of their attacks on wood will be referred to later.
The development of the termites is
apparently simple; the wingless
IO2 The Termites, or White Ants

workers resemble closely, except in size, the just-hatched young; the soldiers
have but to acquire their largeness of head and mandibles, and the perfect
insects their wings. But there is a serious complexity in termite develop-
ment in that at hatching all the young are alike, and the different castes
or kinds of individuals become differentiated during the postembryonic
development, i.e., after hatching. This matter
discussed later. is

In the United States but seven species of this order of insects are known.
They represent three genera, which may be distinguished by the following
table:

KEY TO GENERA.
Simple eyes absent TERMOPSIS.
Simple eyes present.
Tarsi with a pulvillus(little pad) between the claws; prothorax large and
oblong; costal (anterior) area of the wings veined. . CALOTERMES.
Tarsi without terminal pulvillus; prothorax cordate; costal area of wings
without veins TERMES.

Termopsis and Calotermes each include two species, all four limited
to the Pacific Coast; while Termes includes three species, of which but one,
T. flavipes, is found in the northeastern states. This has been introduced from
America into Europe, and is well known there. The other two species, and
flavipes also, are found in the southwestern and Pacific coast states. Thus
Termes flavipes (Figs. 134 and 135) is the only representative of the order Isop-
tera which can be observed and studied in the East,
but it is so commonly distributed that the student of
insects in almost any locality can find its communities.
Despite its abundance, however, the long time it has
been known, and the very interesting nature of its

habits, its life-history is not yet wholly worked out.

FIG. 134. T. flavi- ft makes its nest in or under old logs and stumps.
pes, worker. (Afler ~
oometimes
.

it mines a nest
. .1 i_ r
in the beams and rafters of
jr.
Marlatt; natural
size indicated by old houses. Howard records the serious injuries done
a handsome private residence in Baltimore through
to

the mining of the first-floor timbers by the hidden termites. Comstock

has found them in the southern states infesting living plants, particularly
orange-trees, guava-bushes, and sugar-cane. According to Comstock, they
attack that part of the living plant which is at or just below the surface
of the ground. In the case of pampas-grass the base of the stalk is

hollowed; with woody plants, as orange-trees and guava-bushes, the bark

of the base of the trunk is eaten, and frequently the tree is completely

girdled; with sugar-cane the most serious injury is the destruction of the
seed-cane.
 The Termites, or White Ants 103
The workersof T. flavipes (Fig. 134) are, when full grown, about in.

long, while the soldiers are a little larger. Both of these castes are whitish.
But the winged males (Fig. 1350) and females which come from the nest
and swarm in the air in late spring or early summer are chestnut-brown
to blackish and measure about in. in length. The four wings are of about
equal size, and when the insect is in flight expand about f in. When at
rest they lie lengthwise on the back, projecting beyond the tip of the abdo-
men. They have many veins and are pale brown in color. After flying
some time and to some distance, the insects alight on the ground and shed
their wings (Fig. 13 56). This they are enabled to do because of a curious
suture or line of weakness running across each wing near its base. All the

wing beyond this suture falls off, leaving each now wingless male or female
with four short wing-stumps. These swarming flights
attract the birds. Hagen noted fifteen different species
of birds following such a termite flight one May-day in
Cambridge, Mass. "Besides the common robins, blue-
and sparrows," he says,
birds,
"were others not seen before
near the house. The birds
caught the Termes
partly in
flight, partly on the ground,
and the robins were finally
so gorged in appearance that
"
their bills stood open!
1
After the swarming flight
FIG. I 35 a - FIG. 1356.
the few uneaten males and
females pair, and each pair FIG. 1350. r. flavipes, winged male. (After Mar-
i_ i_i r j , latt; natural size indicated by line.)
probably founds a new colony. Fl(J I3s6 ._ r .
flavipes ,
complementary queen.
Perhaps some of the pairs (After Marlatt; natural size indicated by line.)
are found by workers, and
taken possession of as the royal couple for a new community. Exactly
how the new communities of flavipes begin is not known; and this is
an excellent opportunity for some amateur observer to distinguish himself!
The egg-laying queen mother of a flavipes colony also has yet to be
discovered. There exist in many species of termites individuals called com-
plemental males and females. These are forms which, in case of the loss
of the real king or queea, can develop into substitute royalties. Whether such
forms exist in all flavipes colonies does not seem to be certainly kncwn.
It is obvious that there is still much to learn about the interesting life of
our commonest and most wide-spread termite species.
Of the other six species of our country, all of which are limited to the
southern, southwestern, and Pacific states, three, representing all of the
104 The Termites, or White Ants

three genera, and found about Stanford University,

have been recently
studied by Professor Harold Heath. These are Termopsis angusticollis,
the largest of the American termites, Calotermes castaneus, a small species with
brown-bodied winged forms, and Termes lucifugus, a small white species
common in Europe, and probably brought to this country from there. The fol-
lowing account of Termopsis augusticollis is
based chiefly on Heath's * studies.
Termopsis angusticollis (Fig. 136) is the largest of the species three and
the most abundant. In favorable localities colonies may be found in almost
in dead branches on otherwise healthy
every stump and decaying log, and even
trees. The galleries are made in the deeper portions of the wood, and
usually follow the grain. The colonies with the primary royal pair number
usually from 50 to 1000 individuals, and include workers, soldiers,
and im-
mature forms. The full-grown workers (Fig. 136) are f in. long, the soldiers
a little less, while the
(Fig. 136) f in., and the kings and queen (Fig. 137)
in. After the death of the primary royalties and the
wings expand i^
development of several substitute royal
forms the egg-laying and consequent
increase of the colony are much more
rapid. Heath counted 3221 individuals

FIG. 136. FIG. 137.

FIG. 136. The large termite of California, workers, young,
Termopsis angusticollis;
and a soldier. (From natural size.)
life;
FIG. 137. A, Dealated primary queen of Termopsis angusticollis, at least four years
old; B, complemental queen. (After Heath; three times natural size.)

in one colony, in which were also thousands of eggs. The colony which we
found in the yellow-pine log in the King's River Canon certainly num-
bered many thousands. In the late summer or early autumn the nymphs
(young stage, with visible wing-pads of perfect insects) that have developed
during the year moult, the operation taking from ten to twenty minutes,
after which they rest for two hours, while the wings expand, and the

body- wall hardens and darkens; they take flight usually about dusk. Some
* Heath, H., The Habits of California Termites, Biological Bulletin, vol. 4, 1902,
pp. 47-63.
 The Termites, or White Ants 105

soon fall to the ground, but others may fly a mile. The swarm is pursued
by birds until dark, and then bats take a turn at the chase. The few ter-
mites that escape fly from tree to tree, seeking a spot of decaying wood.
Heath has noted them dashing against door-knobs and nail-holes and against
discolored spots on trees and logs, in their search for a place where decay
has begun. After finding a suitable spot they usually shed their wings,
not by biting them off, as said of some species, but by curving the abdomen
until it rests across the wings of one side and then moving backwards
and sidewise until brought against some obstruction,
the wing tips are
thus causing the wings to buckle and break along the transverse suture or
line of weakness at the base. Sometimes the wings are not shed until after
the nest is begun. The spot is usually selected by the female, and she begins
the mining and does most of it. She is accompanied by one or more males,
who may occasionally help in excavating. When the burrow is large enough
for two,one male usually crowds in beside the queen and fights off the others.
Sometimes two males may remain with the queen; Heath thinks that such
a condition may last for a year or more. He has found a few cases where
two, three, and even six pairs live in company. The actual mating does
not take place, probably, until some time after the nest is begun. Heath
has noted pairing from a week to a fortnight after swarming.
The egg-laying may be Usually, however, about two
long postponed.
weeks and from one to six are deposited
after pairing the first egg is laid,

daily until the total number amounts to from fifteen to thirty. When the
habitat is unusually moist the royal pair may remain together for a year
without producing young. Heath has found the Termopsis royalties to
mate readily in captivity, and has had more than 500 pairs of primary kings
and queens in excellent condition after a year of captivity. Royal pairs
with small colonies are readily found by stripping off the bark of trees from
three to nine months after the swarming period. Heath has been the first
to find actual egg-laying queen termites in this country.
After from fifteen to thirty eggs are laid the laying ceases, and the

parents give their time to enlarging the nest and to caring for the eggs,
which are kept scrupulously clean, and frequently shifted from place to
place in the nest. The young are all alike when first hatched. After three
moults, one of them appears as a large-headed individual, and after three
more moults develops into a perfectly formed soldier, although little more
than one-half the size of the soldiers in old communities. Three months
later another soldier appears, larger than the first, and later others still

larger, until aftera year the full-sized form appears. The first workers,
too, are smaller than the later ones. Nymphs, i.e., young of the winged
individuals, do not appear until after the first year, so that the swarm of
winged individuals cannot leave a nest until the end of the second year of
 The Termites, or White Ants

its existence. The life of these early, undersized individuals is short.

They disappear, perhaps are killed, when the full-sized individuals appear.
These latter, both workers and soldiers, live at least two years and perhaps
longer.
The primary king and queenlive for at least two years, and almost cer-

tainly longer, Heath

believes he has evidence of five years of life. After
the death of the royal pair from natural or other causes, the members of
the orphaned colony develop from the young nymphs from ten to forty sub-
stitute royal forms. By some unknown process, perhaps peculiar feeding,
these selected nymphs are quickly brought to sexual maturity, and the queens

begin egg-laying. As they are fed and cleaned by the workers, their only
business is to lay eggs. Heath observed some of the larger queens to lay
from seven to twelve eggs a day continuously. In exceptional cases a
worker, or even a soldier, may be developed into an egg-laying queen.
One may also occasionally find a few winged soldiers.
In Africa of Termites are known *
forty-nine species and it is (Sjostedt),
on this continent that "the results of Termitid
economy have reached their
climax." More than a century ago an exploring Englishman, Smeathman,
startled zoologists with his account of the marvelous termite communities
of West Africa. He told of the great mound-
nests ofTermes bellicosus, twenty feet high, and
so numerous that they had the appearance of
native villages (Fig. 132). The soldiers are fifteen
times as large as the workers, and the fertile

queen has her abdomen so enlarged and stretched

by the thousands of eggs forming inside that it
comes to be "fifteen hundred or two thousand
times the bulk of the rest of her body and
twenty or thirty thousand times the bulk of a la-
borer." He describes the egg-laying as proceed-
ing at the rate "of sixty a minute, or eighty thou-
sand and upward in one day of twenty-four
hours." In the South Kensington Museum at
London there is such a prodigious queen resem-
bling simply a cylindrical whitish sausage four
FIG. 138. -Worker and
i nc h es long. A similar specimen is to be found
queen of Termes red-
mani. (After Nassonow; in the natural-history museum of the University
natural size.) of Kansas.
The enormous number of individuals in a great village of nests cannot

* Ak.
Sjostedt, Y., Monographic der Termiten Afrikas, Kongl. Svenska, Vetensk.
Handl., v. 34, 1900, pp. 1-236, Stockholm.
 The Termites, or White Ants 107
even be imagined. But according to African travelers the direct results
of the presence of population are very apparent. Drummond
such a

(Tropical Africa, 1891) writes: "You build your house, perhaps, and for
a few months fancy you have pitched upon the one solitary site in the coun-
try where there are no white ants. But one day suddenly the door-post
totters,and lintel and rafters come down together with a crash. You look
at a section of the wrecked timbers, and discover that the whole inside is
eaten clean away. The apparently solid logs of which the rest of the house
is built are now mere cylinders of bark, and through the thickest of them

you could push your little finger. Furniture, tables, chairs, chests of drawers,
everything made of wood, is and in a single night a
inevitably attacked,
strong trunk is and through, and turned into match-
often riddled through
wood. There is no limit, in fact, to the depredation of these insects, and
they will eat books, or leather, or cloth, or anything; and in many parts of
Africa I believe if a man lay down to sleep with a wooden leg it would be
a heap of sawdust in the morning! So much feared is this insect now that
no one in certain parts of India and Africa ever attempts to travel with such
a thing as a wooden trunk. On the Tanganyika plateau I have camped on
ground which was as hard as adamant, and as innocent of white ants appar-
ently as the pavement of St. Paul's, and awakened next morning to find a
stout wooden box almost gnawed to pieces. Leather portmanteaus share
the same fate, and the only substances which seem to defy the marauders
are iron and tin."
But more impressive than this devastation of houses, tables, and boxes is the
sight of millions of trees in some districts plastered over with tubes, galleries,
and chambers of earth due to the amazing toil of the termites in their search
for dead or dying wood for food. According to Drummond, these tunnels
are made of pellets of soil brought from underground, and stuck together
with saliva. The quantity of thus brought above ground is enormous,
soil

and Drummond sees in this phenomenon a result very similar to that accom-
plished by earthworms in other parts of the world, and made familiar to
us by Darwin, namely, a natural tillage of the soil. As Drummond says:
"Instead of an upper crust, moistened to a paste by the autumn rains and
then baked hard as adamant in the sun, and an under soil hermetically sealed
from the air and light, and inaccessible to all the natural manures derived
from the decomposition of organic matters these two layers being eter-
nally fixed in their relations to one another we have a slow and continued
transference of the layers always taking place. Not only to cover their
depredations, but to dispose of the earth excavated from the under-
ground galleries, the termites are constantly transporting the deeper
and exhausted soils to the surface. Thus there is, so to speak, a con-
stant circulation of earth in the tropics, a ploughing and harrowing, not
io8 The Termites, or White Ants

furrow by furrow and clod by clod, but pellet by pellet and grain by
grain."
With a few references to certain special conditions and problems in the
termite economy, we must finish our consideration of these highly inter-
esting insects. Do the termite individuals of a community communicate
with each other, or is the whole life of the colony so inexorably ruled by
instinct that each individual works out its part without personal reference
to any other individual, although with actual reference to all the others,
that is, to the community as a whole ? It is pretty certain that termites have
a means of communication by sounds. The existence of a tympanal audi-
tory organ in the tibiae of the front leg, like that of the crickets and katy-
dids, has been shown by Fritz Miiller, and the individuals have a peculiar

jerking motion which seems likely to be connected with the making of

sounds by stridulation, sounds, however, that are not audible to us.
The spread of termites from one continent to another, as in the case of
Termes flavipes from America to Europe, and Termes lucijugus from
Europe to America, can be easily explained by involuntary migration in
ships. In unpacking several cases of chemicals received from Ger-
many at Stanford University, scores of termites were exposed when the
wooden boxes were broken up. The insects, mining in the wood of the
boxes, had protection, food, and free transportation on their long ocean
journey from Hamburg around Cape Horn to California!
In termite nests are often found individuals of other insect orders. So
often are such cases noted, and so many are the kinds of strangers likely
to be present, that entomologists recognize a special sort of insect economy
which they term termitophily, or love of termites! The strangers seem to
be tolerated by the termites, and apparently live as guests or conmensals.
More than 100 species of insects have been recorded as termitophiles. This
curious guest-life exists on even a much larger scale in the nests of true
ants, in which connection it is called myrmecophily
(see p. 552).
The most important problem, and one whose solution will require much
exact observation (and, if possible, experimentation), is that of the origin,
or causes of production, of the different castes or kinds of individuals in
the termite community. It has been determined by various observers that

all the termites of a community are apparently alike at birth. That is,
there is no apparent distinction of caste, no separation into workers, soldiers,
and perfect insects. The soldiers and workers are not, as was formerly
thought, the result of the arrested development of the reproductive organs.
They are not restricted to one of the sexes. If then it is not arrested develop-
ment, or sex, or embryonic (hereditary) differentiation, what is the causal
factor? Grassi, an Italian student of the termites, thinks that it is food;
that the feeding of the young with food variable in character brings about
 The Termites, or White Ants 109

the differentiation of individuals. To understand this claim it is

necessary
to attend more closely to the feeding habits. The food of termites con-
sistsalmost exclusively, as has already been said, of wood. But this wood
may be taken directly from the walls of the burrow or secured indirectly
from another individual. In this latter case it consists of disjecta of undi-
gested material, which, while mostly wood, must be mixed with other or-
ganic material: because the termites keep their nests clean by eating their
cast skins and the dead bodies of other individuals. This undigested mate-
rial is called proctodeal food. In addition, a certain amount of evidently
very different matter is regurgitated through the mouth from the anterior
part of the alimentary canal. This is called stomodaeal food. As the young
receive all their food from the workers, it is apparent that there is oppor-
tunity for a choice, on the part of the nurses, in the kind of food given the
young. And it is presumed by Grassi that such a choice is made, and that
it results in the differentiation of the castes. As a matter of fact, such a
differentiation of individuals is accomplished in the honey-bee community
by feeding those larvae which the workers wish to make fertile queens "royal
jelly" a rich food regurgitated through the mouth from the anterior part
of the canal.
alimentary This is done for the queens during the whole
larval while larvae which are fed royal jelly for only one or two days,
life,

and then mixed pollen and honey for the rest of larval life, develop into
workers. With the honey-bee, however, the workers are to be looked on
as probably only arrested females. But in the case of Termopsis angusti-
collisHeath has experimented by feeding members of
various colonies, both with and without primary royal

pairs, "on various kinds and amounts of food procto-

deal food dissected from workers, or in other cases from

royal forms, stomodeal food from the same sources,

sawdust to which different nutritious ingredients had
been added but in spite of all I cannot," he says,
"feel perfectly sure that I have influenced in any un-
usual way the growth of a single individual."
This is by all odds the most important and interesting
problem in termite economy, and the solver of it will do
much for zoological science.
A singular and primitive family of small insects, the FIG. 139. Embia
Embiidce, of doubtful affinities, is represented by not texana. (After
more than twenty living species, of which but four Melander; en-
larged.)
occur in this country. The individuals do not live in
communities as the termites do, but in structural characters they probably
more nearly resemble these insects than any others. Fig. 139 illus-
trates a typical Embiid. This species, Embia texana, is about one-quarter
1 1 o The Termites, or White Ants

of an inch long, and of rufous color. It was described from a few specimens
found at Austin, Texas. This insect seems to be very susceptible to differ-
ing degrees of humidity, and specimens were visible only after the ground
had been moistened by rains. As the sun dries the ground, the insects
turrow into the soil. They spin small silken webs in which they live singly.
These webs are tunnels made in some crevice of the rock which shelters
them, or are spun between grains of soil. They are an inch or more in
length and closed at one end, and probably serve simply for protection. The
spinning-organs of the insect are located in its fore feet, a condition unique
among animals. The food-habits of the Embiids are not yet known.
 CHAPTER VIII

THE BOOK-LICE AND BARK-LICE (Order Corrodentia)

AND THE BITING BIRD-LICE (Order Mallophaga)
OMETIMES in taking from the shelves an old
book, long untouched, there may be seen, on
turning its leaves, numerous extremely minute,
pale-colored, wingless insects, the book-lice, or
dust-lice. So small are they, indeed, that a
reading-glass or hand-lens will be needed to make
out anything of their real appearance. They
run about rather swiftly and seek to conceal their soft, defenceless little

bodies somewhere in the binding. Under the lens they are seen to have
a rather broad, flattened body (Fig. 140), six short legs, no wings (although
sometimes tiny wing-pads are present), long, slender antennae, and a pair of
small black spots on the head, the simple eyes. There is a distinct neck,
the head being free, and plainly wider than the prothorax. The abdomen
is nearly oval in outline. There are no distinctive markings or pronounced
chitinization of the
soft body-wall. These book-lice can be found else-
where than old books; they feed on dry, dead organic matter, the
in

paste of the book-bindings and the paper, and are common in birds' nests,
where they feed on the cast-off feathers, in the crevices of bark, and on
old splintered fences, where they feed on moulds and dead lichens.
Certain other insects closely related to the book-lice are not so small and
simple, however, some having two pairs of wings and a plump, rounded
body (Fig. 141); these look much like plant-lice (Aphids). These winged
kinds do not live in libraries, moreover, and the name "book-lice" is a
misnomer for them. They are rarely seen by persons not trained entomolo-
gists, and indeed are not at
all familiar to professed students of insects.

The life-history of these obscure insects has been but little studied, but it

is a simple kind, the metamorphosis being incomplete, and in the case

of
of the wingless forms certainly very slight. The young of the wingless forms
"
greatly resemble the old, but have no ocelli or wings, and sometimes the
tarsi are of two joints, while in the adult they have three." The structure
of the adults presents no points of particular interest except in the case of
the mouth. The book-lice have biting mouth-parts, the jaws being strong
and heavy for the successful mastication of the hard dry food. In the throat
1 1 2 Book-lice and Bark-lice ; Biting Bird-lice

there is a peculiar little chitinized structure, which may be called the

oesophageal sclerite (Fig. 145). This structure is situated in the floor of

the pharynx (forward end of the oesophagus), and has some special function
in connection with the peculiar food-habits. It was first described by Bur-

gess, and was for a long time supposed to be peculiar

to the book-lice alone. But, in a study of the mouth
structure of the biting bird-lice (Mallophaga) I found ,

an almost identical cesophageal sclerite in thirteen out

of the twenty-two genera of the Mallophaga. On
the basis of this common possession of a curious
and undoubtedly important mouth structure by the
book-lice and the bird-lice (and on the basis of other
strong similarities) it seems certain that these two
groups of insects have a common ancestry not very
remote, and probably should be included in a single
FIG. 140. A wingless order.
ouse k P* Atr S P-
The order Corrodentia as at present known con-
K? l
(Much enlarged.)
,'

tains about two hundred described species, scattered

over the world. The largest species occur in Brazil, and have an ex-
panse of wing of nearly an inch. Ceylon and the Hawaiian Islands are
said by Sharp to be specially rich in species.
The members of the order can be divided into two families as follows:

Wings well developed; ocelli present (in addition to compound eyes) ___

Wings wanting or present as small scales or pads; ocelli absent ____ ATROPID^E.

The winged Corrodentia or Psocidae (which may be called bark-lice to

distinguish them from the wingless book-lice) are
too rarely seen to be at all familiar. They may
most commonly be found in small clusters on bark,
each cluster or colony being covered over by fine
silken threads spun from the mouth. The wings
are held roof-shape over the back (Fig. 141), and
the body and wings are usually pale smoky in FlG I4 j A _ w l nge d
general color. The small white eggs are laid on the bark-louse. (Thirteen
size -) times natural
surface of the bark in small patches, and in a cluster
of bark-lice, individual, in all stages, from very young to adult, may be seen.
Banks gives the following key to the North American genera:
The techinal terms discoidal cell and posterior cell may be understood by reference to
Fig. 142.
1. Wings with scales and long hairs ......................... AMPHIENTOMUM.
Wings without hairs and scales, hyaline ..................................... 2.
2. Tarsi 3-jointed ......................................................... 3.
Tarsi 2-jointed ......................................................... 4.
 Book-lice and Bark-lice ; Biting Bird-lice 1 1
3

3. Discoidal cell closed MYOPSOCUS.

Discoidal cell open ELIPSOCUS.
4. Discoidal cell closed 5.
Discoidal cell open 6. 2
^\ 5
(

5. Discoidal cell four-sided. Psocus. : ^^i~^Z~-~~^ *S^

Discoidal cell five-sided.
AMPHIGERONTIA.
6. Third posterior cell elliptical.

C.ECILIUS.
Third posterior cell elongated.
POLYPSOCUS ^ IG- J 4 2 Diagram of venation of a Psocid.
'

d, discoidal cell; ia. za, -la, posterior cells.

Third posterior cell absent.
^ fter
( Banks-)
PERIPSOCUS.

The few North American species of the true book-lice or Atropidae are
included in five genera, which may be distinguished as follows:
The technical terms, hitherto undefined, used in the following table are the following:
squama, wings in the condition of small scales or pads; hyaline, clear, not colored.
1. Meso- and metathorax united, no wings ATROPOS.
Meso- and metathorax separate, rudimentary wings 2.

2. Wings with veins DORYPTERYX.

Wings veinless, in form of squamae or tubercles 3.

3. Squamae small, hyaline CLOTHILLA.

Squamae in the form of scars LEPINOTUS.
Small tubercles in the place of squamae HYPERETES.

The genera Atropos and Clothilla were named for two of the three Fates
of mythology, and a third genus was named Lachesis for the third Fate, but
unfortunately the last genus was not a valid one, so the book-lice have lost
their third Fate, and by the rigid laws of zoological nomenclature can never

regain her! The few species of these two Fate-named genera are the com-
monest of the book-lice. Atropos divinatoria is the species usually
found in books. It is about i mm. long, is grayish-white, and the small
eyes show as distinct black specks on the head. It does not limit its feeding
to the paste of book-bindings, but does much damage to dried insects in
collections. To has long been attributed the power of producing
this insect
a ticking noise known as the "death-watch," but McLachlan, an authority
on the Corrodentia, does not believe that this minute insect "with a body
so soft that the least touch annihilates it can in any way produce a noise
sensible to human A
small beetle, called Anobium, is well known
ears."
to make such a ticking (by knocking its head against the wood of door-casings,

floors, etc., in which it lives) and probably the "death-watch" is always

made by this beetle.

BIRD-COLLECTORS are often annoyed by small, wingless, flat-bodied, swift-

running insects which sometimes escape from the feathers of bird specimens
to the hands and arms of the collector. Poultry-raisers are sometimes more
seriously troubled by finding them so abundant on their fowls as to do con-
114 Book-lice and Bark-lice; Biting Bird-lice

siderable injury. They are called bird-lice, but they should not be confused,
because of this name, with the true blood-sucking lice that infest many kinds
of animals, particularly domestic mammals and
uncleanly persons. The
biting bird-lice(Fig. 143), constituting the order Mallophaga, never suck
blood, but feed exclusively on bits of the dry feathers, which they bite off
with small but strong and sharp-edged mandibles. The true lice have
mouth-parts fitted for piercing and sucking, and
constitute one of the numerous families of the
order of sucking bugs, Hemiptera (see p. 217).
More than a thousand species of biting bird-
lice,or Mallophaga, are known, of which about
two hundred and fifty have been found on North
American birds. Although by far the larger num-
ber of Mallophaga infest birds, numerous species
are found on mammals. On these hosts the insect
feeds on the hair or on epidermal scales. On
both birds and mammals, therefore, the food con-
sists of dry and nearly or quite dead cuticular sub-
stances, and never of blood or live flesh. Those
species of Mallophaga which infest birds are never
found on mammals, and vice versa.
The in J UI7 done to the hosts b X thes e parasites
consists not in the character of the food-habits, but
chiefty in the irritation of the skin caused by the

George E. Mitchell; scratching of the sharp-clawed little feet of the insects

naturalsize.one-twelfth in their migrations over the body. When, as hap-
mch.) \
sometimes in poultry-yards and dovecotes,
pens
a fowl or pigeon is infested by hundreds of these active little pests, the
afflicted bird becomes so restless and excited that it takes too little food

and gets too little rest and thus grows thin and weak. The dust-baths
taken by fowls and other birds are chiefly to get rid of the bird-lice. The
fine dust, getting into the breathing-pores (spiracles) of the insects, suffocates
them. So that the best remedies for these pests of the barn-yard are to
see thatthe fowls have plenty of dust to bathe in, and also to keep

thoroughly clean their roosting- and breeding-places. By tightly closing

up the hen-house and burning sulphur inside (the fowls, it is hardly necessary
to say, first being excluded) most of the infesting parasites can be killed.
Thelife-history of the Mallophaga is very simple. The small elongate
eggs are glued separately to the hair or feathers of the host, and from them
young soon hatch (Fig. 144,3), which, except in size and, to some degree, in
marking, closely resemble the parents. These young begin immediately their
hair or feather diet, grow larger, moult a few times, and in a few weeks reach
 Book-lice and Bark-lice ; Biting Bird-lice 115

maturity. There is never, in young or old, any sign of wings or wing-pads.

The body is flattened, so much so indeed that it is difficult to hold a live

specimen securely between thumb and finger-tip. The body-wall is strongly

chitinized, and is firm and smooth. The markings are often very distinct,
and sometimes bizarre, but the coloration varies only from white to black
through various shades of pale yellowish brown, tawny, reddish brown, and
blackish brown. The antennae are short and in one suborder (see classifica-
tion key) are wholly concealed in pits or grooves on the under side of the

FIG. 144. Immature and adult stages of the biting bird-louse, Lipeurus forficulatus,
taken from a pelican, adult female; 2, adult male; 3, very young stage; 4,
i,
older immature stage. (Natural size of adult specimens in.)^
flattened head. The legs are strong, and each foot bears two claws. These
small creatures run very swiftly.
Perhaps the oddest thing about the structure of the Mallophaga is the
presence in the throat of the curious cesophageal or pharyngeal sclerite
already referred to in the account of the Corrodentia. This sclerite is a
sort of bonnet-shaped piece (Fig. 145) lying in the lower wall of the throat
and seems to be an arrangement for starting the little bitten-off pieces of
feather barbs straight, that lengthwise down the oesophagus! The bark-
is,

lice and book-lice, which have a similar cesophageal sclerite, also bite off

and swallow small bits of hard, dry organic substance.

 1 1 6 Book-lice and Bark-lice ; Biting Bird-lice

The most interesting thing in connection with the Mallophaga, excepting

their parasitic life and strange food-habits, is the puzzling problem of their
distribution. The problem in its largest phase is this: the species of Mal-
lophaga are, in a majority of cases, peculiar (so far as recorded) each to some
one host species. But the instances are many where a single parasite species
is common to a few or even to many host species. How does this condition
of commonness to several hosts come to exist?
As the Mallophaga are wingless, their power of migration from bird to
bird is limited. Moreover, they can live for but a short time off the body
of a warm-blooded host. After a bird is
shot, the Mallophaga on it die in from
two hours to three or four days: in rare
cases living individuals are found on the
drying bird-skin after a week. Although
the parasites in a badly infested hen-house
willbe seen on the roosts and in the nests,
in Nature the insects are rarely found off
the host's body. On such a likely place
as an ocean rock from which I had just
frightened hundreds of perching pelicans,
cormorants, and gulls no parasites could
be found. Practically migration must be
accomplished while the bodies of the hosts
FlG. 145. Bonnet-shaped pharyngeal ar( c 1]r u rac pc
mntart
ntact '
'
, .

from a biting
sclerite, lateral aspect,
bird-louse, Eurymo$tus taurus, from mating and nesting, and when gregarious
an albatross. (Greatly magnified.) b j rds roost Qr
perch dosely together-
Occasional migration might occur from a bird of prey to its captured
victim, or from victim to hawk.
The general character of the cases in which a single Mallophagan species
is common to several host-species may now be considered. Docophonis
larihas been found on thirteen species of sea-gulls, and Nirmus lineolatus
on nine. Gulls are gregarious, perching together on large food-masses
and on ocean rocks. But on these rocks gulls are closely associated with
other coast birds, as cormorants, pelicans, murres, etc. And the gull-para-
sites might have opportunities to migrate to these other bird-species.
Docophorus icterodes and Trinoton luridum are common to many duck species
(each has been collected from nine), but ducks also are gregarious, and in
addition are much given to hybridizing. But a parasite may be common to
several host-species of non-gregarious habits. Docophorus platystomus is
common to several hawk-species, D. cursor to several owl-species, D. excisus
to several swallows, D. californiensis to several woodpeckers, and D. com-
munis to several passerine birds. In the other genera of Mallophaga are
 Book-lice and Bark-lice ; Biting Bird-lice 1 1
7

similar cases, and in all these cases it is hard to see how actual migration
of the parasitefrom host to host of different species could take place. Indeed
there are cases in which such migration is absolutely impossible. Of the
262 species of Mallophaga taken from North American birds, 157 have
been described as new species, while 105 are specifically identical with Mal-
lophaga originally described from European and Asiatic birds; hosts, that
is, not only of different species, but geographically widely separated from
the North American hosts! Eliminating the few cases of importations of
living European birds to this country, and the few species of cicumpolar
range, there remain to be accounted for about 100 cases in which a single
species of Mallophaga is common to both Old World and New World hosts.
It will have been noted that in all the cases above mentioned of parasite
species common to several North American host-species, the host-birds are

closely allied forms, that is, species of the same genus or allied genera.

This condition holds good also which both

for practically all of the cases in

European and American hosts have a common parasite. For example,

Docophorus pertusus is common to the European coot (Fulica atra) and
the American coot (Fulica americana) Nirmus pileus is common to the
;

European avocet (Recurvirostra avocetta), and to the American avocet

(Recurvirostra americana) ; Lipeurus jorficulatus is common to the European
pelican (Pelecanus onocrotalus) and to the American pelicans (Pelecanus
erythrorhynchus and P. calijornicus), and so on through the list. From
this fact of near relationship of hosts in all the cases of parasite species com-
mon to several host-species it seems almost certain that this common occur-

rence, under circumstances not admitting of migration of the parasites from

host to host, is due to the persistence of the parasite species unchanged from
the time of the common ancestor of the two or more now distinct but closely
allied bird-species. In ancient times geographical races arose within the limits
of the ancestral host-species these races or varieties have now come to be dis-
;

tinct by superficial differences in color and mark-

species, distinguished
ings of plumage, etc. But the parasites of the ancient hosts have remained
unchanged; the plumage as food, the temperature of the body, practically
the whole environment of the insect, have remained the same; there has
been no external factor at work tending to modify the parasite species, and
it exists to-day in its ancient form, common to the newly arisen descendants

of the ancient host.

To classify Mallophaga the following keys to suborders, families, and
genera may In these keys are included only genera which have
be used.
been found in the United States. Seven other genera of Mallophaga are
known.
In the following tables the following technical terms are used which have not been
previously defined: cla-vate, club-shaped; capitate, with the tip swollen like a ball; tra-
1 1 8 Book-lice and Bark-lice ; Biting Bird-lice

becula, triangular membranous processes projecting laterally from the head and situated
one in front of each antenna; temples, the hinder lateral parts of the head; ocular emar-
gination, a bending in of the lateral margins of the head just in front of the eyes; labral
lobes, short blunt membranous processes projecting laterally from near the front angles
of the head; sternal markings, blackish markings, bars or spots, on the ventral aspect of
the thorax.
KEY TO SUBORDERS OF MALLOPHAGA.
With short slender 3- or 5 -segmented, exposed antennae; no palpi; mandibles
vertical ISCHNOCERA.
With short clavate or capitate, 4-segmented antennae concealed in shallow cavities
on under side of head; 4-segmented palpi; mandibles horizontal.
AMBLYCERA.
KEY TO GENERA OF THE SUBORDER ISCHNOCERA.
A. With 3-segmented antennas; tarsi with one claw; infesting mammals only (family
TrichodectidcE) TRICHODECTES.
AA. With 5 -segmented antennae; tarsi with two claws; infesting birds only (family
Philopterida) .

B. Antennae alike in both sexes.

C. Front deeply angularly notched AKIDOPROCTUS.
CC. Front convex, truncate, and rarely with a curving emargination, but never
angularly notched.
D. Body broad and short; head with large movable trabeculae.
E. Forehead with a broad, transverse membranous flap, project-
ing beyond lateral margins of the head in the male, barely
projecting in the female GIEBELIA.
EE. Without such membranous flap DOCOPHORUS.
DD. Body elongate, narrow; head with very small or no trabeculae.
NIRMUS.
BB. Antennae differing in the two sexes.
C. Body wide, elongate oval to suborbicular.
D. Temples rounded; tip of abdomen with shallow, curving emargina-
tion; antennae with no appendage; third segment unusually long.
EURYMETOPUS.
DD. Temples usually angulated; tip of abdomen convex, rarely angularly
emarginated with two points.
E. First antennal segment of male large, and sometimes with
an appendage; segment always with appendage.
third
GONIODES.
EE. First antennal segment of male large, but always without
appendage; third segment without appendage. .GONIOCOTES.
CC. Body elongate, narrow, sides subparallel.
D. Antennae and legs long; a semicircular depression in front of
mouth LIPEURUS.
DD. Antennas and legs short; depression in front of mouth narrow and
elongate, extending as a furrow to the anterior margin of the head.
ONCOPHORUS.
KEY TO GENERA OF THE SUBORDER AMBLYCERA.
A. Tarsi with one claw; infesting mammals only (family Gyropidce) GYROPUS.
AA. Tarsi with two claws; infesting birds only (family Liotheidce).
B. Ocular emargination distinct, more or less deep.
 Book-lice and Bark-lice ; Biting Bird-lice 119
C. Forehead evenly rounded, without lateral swellings; antennae projecting
slightly beyond border of the head COLPOCEPHALUM.
CC. Forehead with strong lateral swellings.
D. Mesothorax separated from metathorax by a suture TRINOTON.
DD. Meso- and metathorax fused; no suture L^MOBOTHRIUM.
BB. Ocular emargination absent or very slight.
C. Sides of the head straight or slightly concave, with two small laterally
projecting labral lobes PHYSOSTOMUM.
CC. Sides of the head sinuous; forehead without labral lobes.
D. Ocular emargination filled by a strong swelling; sternal markings
forming a quadrilateral without median blotches NITZSCHIA.
DD. Ocular emargination without swelling, hardly apparent or entirely
lacking; median blotches on sternum.
E. Very large; with two-pointed appendages on ventral aspect
of hind head; anterior coxae with very long lobe-like append-

ages ANCISTRONA.
EE. Small or medium; without bi-partite appendages of hind head.
MENOPON.

The Mallophaga most likely to come under the observation of people

not collectors of birds are the species which infest domestic fowls and mam-
mals, and the following few descriptions and figures of particular species
are therefore limited to such kinds.
The most notorious member of the order is the common chicken-louse,
Menopon pallidum (Fig. 146). It is of a pale straw-yellow color, from
i mm. l
(% j in.) to 1.5 mm.
in length, and is an un-
usually swift and active
little pest. Other Mallo-
phaga infesting chickens
are Goniocotes hologaster,

recognized by its squarish

head with angulated
temples, and Lipeiirus
variabilis, 2 mm. (-fa in.)

long and slender, with dis-

tinct black markings on
the otherwise smooth,
white body.
Ducks are infested by
several species. Com- F _,

mon among them is the FIG. 146. The biting chicken-louse, Menopon pallidum.
little DocophorilS icterodes (After Piaget; natural size, i to 1.5 mm.)
FIG. 147. The biting louse of wild ducks, Docophorus
/T-,.
fig-M?;*
N
1 m -
f
\T5
.,
m \
-) icterodes. (Natural size indicated by line.)
long, with head curiously
expanded and rounded in front, darkish-red head, and thorax with darker
I2O Book-lice and Bark-lice; Biting Bird-lice

bands, and a white region in the middle of the abdomen. Trinoton luridum
is another common duck-louse unusually large, being from 4 to 5 mm. ( 5 in.)

long and readily distinguished by the triangular

head with lateral swellings, and the abdomen with
pronounced blackish-brown transverse bands.

FIG. 150.

FIG. 152.

FIG. 148. A biting louse of pigeons, Lipeurus baculus. (Natural size indicated by line.)
FIG. 149. Biting louse of the dog, Trichodectes latus. (After Nitzsch; natural size,
i to 1.5 mm.)
FIG. 150. Biting louse of the horse, Trichodectes parumpilosus, male. (After Morse;
natural size shown by line.)
FIG. 151. Biting louse of cattle, Trichodectes scalaris. (After Lugger; natural size, 1.5
to 2 mm.)
FIG. 152. Biting louse of fringilline birds, Docophorus communis. (Natural size in-
dicated by line.)
 Book-lice and Bark-lice; Biting Bird-lice 121

Pigeons are almost always infested by a long and very slender louse,
Lipeurus baculus (Fig. 148). The head and thorax are reddish brown,
while the abdomen is dusky with darker segmental blotches. This bird-
louse was described and named more than two hundred years ago.
All of the species infesting domestic mammals belong to the genus Tricho-
dectes. Dogs are often infested by Trichodectes latus (Fig. 149), a short,
wide-bodied species about i mm. long; while cats are less often infested by
T. subrostratus, distinguishable by the rather pointed head with a short,

longitudinal furrow on the under side. Horses and donkeys are troubled
by two or three species, of which T. pilosus, a hairy form with antennae rising
near the front of the head, and T. parumpilosus (Fig. 150), a broader-bodied
form with head larger and less flatly rounded in front, are most common.
Trichodectes scalaris (Fig. 151) infests cattle the world over, while sheep
and goats have species peculiar to themselves. Comparatively few species
of Trichodectes have been recorded
from wild mammals, but this is
simply because they have not been
sought with care. Species have

FIG. 153. FIG. 154.

FIG. 153. A biting louse of gulls, Nirmus felix, male. (Natural size indicated by line.)
FIG. 154. Giant bird-louse of the albatrosses, Ancistrona gigas, male. (Natural size
indicated by line.)

been found on the bear, raccoon, fox, coyote, weasel, gopher, beaver, deer,
skunk, and porcupine. Gyropus, the other mammal-infesting genus of
122 Book-lice and Bark-lice; Biting Bird-lice

Mallophaga, has been found only on the guinea-pig. Washing the body of
the infested animal with kerosene emulsion (see p. 190) is probably the
most effective remedy for biting lice.
Of the nearly three hundred species of Mallophaga which I have recorded
(Proc. Nat. Mus., v. 22, 1899, PP- 39~ IO ) from wild North American birds,
mention may be made of the largest, Lcemobothnum loomis, taken from the
Canada goose; Docophorus communis (Fig. 152), the most abundant and
of

widely distributed parasite of perching and song birds; of the pretty Nirmus
jelix (Fig. 153), with its clean white body and sharply marked black spots;
of the fierce-looking Lipeurus ferox, found on albatrosses; and of Ancistrona

gigas (Fig. 154), found on fulmars, the broadest of the Mallophaga.

As there are nearly one thousand different species of North American
birds, and Mallophaga have been taken from but two hundred and fifty of
them, it is obvious that the collector and student of these parasites has a

profitable field open to him.

 CHAPTER IX

THE COCKROACHES, CRICKETS, LOCUSTS, GRASS-

HOPPERS, AND KATYDIDS
(Order Orthoptera)
1

E do not shut up our singing insects in cages

as the Japanese do, and bring them into
the house to cheer or amuse us, but we do

enjoy them, and were our summer and

early fall days and nights to become sud-
denly silent of chirping and shrilling, we
should realize keenly how companionable
crickets and grasshoppers and katydids
had been for us. A wholesome blitheness
and vigor and ecstasy of living rings out
in the swift and steadfast song of most of
our field and wood insect singers, while
the cheeriness of the cricket on the hearth
is familiar poetryand proverb.
Almost insect music comes from the members of one order, the
all this

Orthoptera. Indeed there is but one famous insect maestro, the cicada (of the
order Hemiptera), which does not belong to the group of crickets, locusts, green
grasshoppers, and katydids. Besides being singers, too, the Orthoptera
are the characteristic leapers of the insect world; crickets and locusts easily

surpass the world's athletes for high jumping if the record takes into account
the comparative size of the athletes. And, curiously, the singing Orthoptera
are the leaping ones. Of the six families composing the order, three include
insects which do not sing nor leap, while the other three are made up of
singers and leapers.
As one tramps the roadways or dry pastures in summer and autumn,
the steady shrilling of the locusts on the ground, or their sharp "clacking"
as they spring into air, are most familiar sounds. When you ramble through
the uncut meadows and lush low grounds the still shriller singing of the
slender-bodied, thin-legged, meadow green grasshopper is heard, while
in the orchards and woods the snowy tree-crickets and broad-winged katydids
123
i
24 Cockroaches, Locusts, Grasshoppers, and Crickets

keep up the chorus. At home, in house and garden, the domestic cricket
music to the already over- full ears. All this choiring is done by
offers its

singers without a voice; that is, without the

production of sound from the throat and
mouth by means of vocal cords set into vi-
bration by air.Insects are orchestral per-
formers, using their legs and wings, for the
most part, to make their music. When the lo-
cust sings while at rest, it is rasping the inner
surface of the broad hind thighs across the

roughened outer surface of the folded fore

wings; when it "clacks" in the air, it is strik-
ing the front margin of the hind wing back
and forth past the hinder margin of the
thickened fore wings. When the cricket
shrills on the hearth, or anywhere else, he, for
only the male crickets have the musical gift, is holding
_^_ n * s ^ore W ' n 8 s U P over hi s body at an angle with it of
VI X. Vlll
1
^xJl about 45 and is rubbing together the upper surfaces of
x\J
the basal region of the fore wings, which are specially
modified for this purpose. The tree-crickets, katydids,
and meadow green grasshoppers have, in the males,
the same general sort of music-making apparatus as
the cricket, and sing by similarly rasping or rubbing

together the modified parts of the fore wings. This

FIG. 155. Longitudinal section through head and neck of locust,

showing disposition of alimentary canal, brain, and sub-
cesophageal ganglion. (After Snodgrass; much enlarged.)

music-making by rasping is called stridulation, and for the most part

insect stridulation is strictly strident, and sounds to better advantage in the
field than it would from caged songsters in the parlor.
 Cockroaches, Locusts, Grasshoppers, and Crickets 125

All the Orthoptera have biting mouth-parts, and bite off and chew their

food, which is usually live

vegetable matter,
especially green leaves,
although the members of one family are predaceous, preying on other insects,
and those of another family prefer dried vegetable or animal matter. The
metamorphosis incomplete, the young, when hatched, resembling the parents
is

and lack of wings. The young have the same feeding

except for small size
habits and same haunts as the adults, and by development and growth the

FIG. 156. The immature stages of a locust, Melanoplus femur-rubrum. a, just hatched,
without wing-pads; b, c, d, and e after first, second, third, and fourth moultings
respectively, showing appearance and development of wings; /, adult, with fully
developed wings. (After Emerton.)

wings and parental stature are soon acquired. The name of the order is
derived from the straight-margined leathery fore wings, or elytra, whose
chief function is to cover and protect the larger membranous hind wings
on which the flight function depends. Among the leaping Orthoptera the
hindmost legs are very large and long, and when at rest or in walking the
"knee-joints" of these legs are much higher than the back of the insect.
The three singing and leaping families are the Acridiidae, locusts and

grasshoppers with short antennae; Locustidse, meadow green grasshoppers

 126 Cockroaches, Locusts, Grasshoppers, and Crickets

and katydids, all with long thread-like antenna?; and Gryllidae, the crickets.
The three silent and walking or running families are the Blattidae, cock-
roaches; Mantidae, praying-horses and soothsayers; and Phasmidae, walk-
ing-sticks or twig-insects. These families can be distinguished by the follow-
ing table:

KEY TO FAMILIES OF ORTHOPTERA.

Non-leaping and mute; hind femora closely resembling those of the other legs and
scarcely stouter or longer than the middle femora; tarsi 5 -segmented; ovipositor
concealed.
Body oval, depressed; head nearly horizontal and nearly or quite concealed by
the flattish shield-like pronotum; quickly running. . . .(Cockroaches.) BLATTID^E.
Body elongate, generally narrow; head free, often with constricted neck; pronotum
elongate, never transverse; slowly walking.
Fore legs spined and fitted and held for grasping; antennae usually shorter than
body; pronotum usually longer than any other body segment; anal cerci
jointed (Praying Mantes.) MANTID^:.
Fore legs not fitted for grasping; antennae usually longer than body; pronotum
short (Leaf-insects and Walking-sticks.) PHASMID^E.
Leaping and usually capable of stridulation; hind femora stouter or longer, or both,
than the other femora; the hind legs enlarged, for leaping; tarsi 4- or 3-segmented;
head vertical; ovipositor usually visible.
Antennae much shorter than the body (with few exceptions) ocelli three tarsi 3-seg-
; ;

mented; auditory organs, when present, situated on basal abdominal segment;

ovipositor composed of two pairs of short, strong, slightly curving pieces.
(Locusts.) ACRIDIID.E.
Antennas much longer than the body, delicately tapering; tarsi 3- or 4-segmented;
auditory organs usually near the base of the fore tibiae; ovipositor usually pro-
longed into a compressed blade, or needle, its parts compact.
Tarsi 4-segmented; ocelli usually absent; ovipositor usually exserted and forming
a strongly compressed, usually curving, blade with tip not expanded.
(The long-horned grasshoppers.) LOCUSTID^E.
Tarsi 3-segmented; ocelli variable; ovipositor usually exserted and forming a
nearly cylindrical straight needle, the tip somewhat expanded.
(Crickets.) GRYLLID^E.

Mrs. Smith takes itamiss when you ask permission to collect "roaches"
in her house, and will prove to you any day the conspicuous absence of these
unwelcome guests in the scrubbed and spotless pantry and kitchen. But
with a candle go stocking-footed at night into the same kitchen and you
will not unlikely find "good hunting." Although but few of the thousand
different kinds of cockroaches known in the world are to be found in the

United States, these few, and particularly three or four imported foreigners
among them, are very abundant, and, after dark, very much in evidence in

their favorite habitat. Their chosen abiding-place is in kitchens, pantries,

laundries, restaurants, bakeshops, etc., where the atmosphere is warm
 Cockroaches, Locusts, Grasshoppers, and Crickets 1
27
and humid and the roach's table is well set with good things. Almost any
sort of dry organic matter suits their taste; bread, crackers, miscellaneous
cold-lunch delicacies, the paste of bookbindings and wall-paper, leather,
woolens, and even their own egg-cases and cast skins making up the dietary.
The fo'k'sel and galley of ships are the roaches' special joy; the hotels and

restaurants of tropic and subtropic lands house swarms of these bill-evading

guests. From Mazatlan, Mexico, a naturalist sent me quarts of large native

American roaches (Periplaneta americana), which he readily scooped up
from his bedroom floor. Ships come into San Francisco from their long
half-year voyages around the Horn with the sailors wearing gloves on their
hands when asleep in their bunks in a desperate effort to save their finger-
nails from being gnawed off by the hordes of roaches which infest the
whole ship. A few of our species still live outside under stones and old
logs, but most of them have learned that an easier life awaits them in the
kitchen.
The roaches compose the Orthopterous family Blattidae, and are an
ancient and persistent insect group. In Carboniferous times, before flies,
butterflies, bees, and wasps had come into existence, cockroaches were
the dominant insects. The body in all is flattened and slippery with the
legs adapted for quick running, so that the insects are well fitted to escape
safely into narrow cracks and crevices. The head is concealed from above
by the expanded shield-shaped dorsal wall of the prothorax (pronotum).
Wings are present in most species, the front pair
leathery and serving, when the wings are folded, to
cover and protect the larger, thin, membranous
hind pair. In some forms the females are wingless,
and the indoor habit may be held responsible for
the lessened usefulness and resultant loss of the FlG r 57-
gg-case of -

mi ^i ^^ j r L-.L- i j cockroach. (Three times

wings. The mouth-parts are ntted hard for biting natural size.)
dry substances, the jaws being strong and toothed.
The eggs are laid in small purse-like, horny, brown cases (Fig. 157), which
are usually carried about by the female until the young are ready to issue.
The young grow slowly, requiring probably about a year, in most species,
tobecome fully developed. From the beginning, the young can run about
and take care of themselves, eating the same kind of food as the adults.
They moult several times during growth, and at each moult the wing-pads
are a little larger.

There are four common species of cockroaches found in dwellings in this

country, only one of which is native. This is the large American roach,
Periplaneta americana, about i inches long (to tip of folded wings), light
brown in color, and with the wings expanding nearly 3 inches. This species
is abundant in the middle and western states, having gradually extended
 128 Cockroaches, Locusts, Grasshoppers, and Crickets

its range north from its native region in Mexico and Central America. The
Australian roach, Periplaneta australasia, resembles P. americana, but is
darker in ground color, a quarter of an inch shorter, and has a conspicuous
yellow submarginal band running around the shield-shaped pronotum.
Each wing has also a strong yellow tapering bar in the basal part of
fore
the costal region. It came originally from the Australian Pacific
region,
and is now spread widely over the world, being common in this country
in Florida and other southern states. The most abundant and destruc-
tive house-roach in the eastern states is the small German cockroach,

Ectobia germanica (Fig. 158), about half an inch long, and pale yellowish
brownish with a pair of distinct black longitudinal stripes on the pro-

FIG. 158. FIG. 159. FIG. 160.

FIG. 158. The croton-bug, or German cockroach, Ectobia germanica. (Twice natural
size.)
FIG. 159. The black beetle, or Oriental cockroach, Periplaneta orientalis. (One and
one-half times natural size.)
FIG. 1 60. The common wood cockroach, Ischnoptera pennsylvanica. (After Lugger;
natural size indicated by line.)

notum. This roach is often called croton-bug, from its intimate asso-
ciation with the pipes of New York City's Croton-water system. It is an

importation from Europe, where it is especially abundant in Germany. Its

real nativity is unknown, but it is now of world-wide distribution. The
fourth species is the black or Asiatic roach, or black beetle, as sometimes
it is

called, Periplaneta orientalis (Fig. 159). This roach is about one inch
long, with brownish-black body; in the female the wings are rudimentary,
and in the male the wings when folded do not quite reach the tip of the
abdomen. This species is common in all the eastern and Mississippi
Valley states and extends west as the great piains. It is the
as far
commonest cockroach in England and Europe. The native outdoor species
most familiar in this country is the common wood-cockroach, Ischnoptera
 Cockroaches, Locusts, Grasshoppers, and Crickets 1
29

pennsylvanica (Fig. 160), with long, light-colored wing-covers, and wings

which extend considerably beyond the tips of the abdomen. The margin
of the pronotum is light, the disk being dark, and the front margins

(lateral when folded) of the wing-covers are lighter than the discal
parts. The body is an inch long and rather narrow and slender. This
is common in the woods and sometimes comes into houses in
species
summer-time.
In the southern states and those of the Mississippi Valley a large insect
may be not infrequently seen standing motionless in a corner of a window,
in a striking attitude. This attitude may be taken as one of hopeful prayer,
as those who gave the name praying-mantis to the insect seem to have taken
it, or one
of self-confident readiness to do violent work with those upraised,

sharply spined, and very willing fore feet. This is the way the house-flies

rightfully
take the mantis's attitude. Watch an unwary bluebottle crawl or
buzz into the fatal corner. Blundering buzziness is finished for that blue-

FlG. 161. The praying-mantis, Mantis religiosa. (After Slingerland; natural size.)

bottle; and the first course of a square meal has come to him who waits
and watches. Other names, as rearhorse, camel-cricket, and soothsayer,
have been given the mantis, all suggested by the attitude and curious body
make-up of the creature. The prothorax is long and stem-like, the head
broader than long, with protuberant eyes, and the fore legs are not used
for locomotion, but are large, strongly spined, and fitted for seizing and hold-

ing the prey. The wings are short and broad and usually rather leaf-like
in coloration and texture, the whole insect when at rest resembling somewhat
a part of the plant on which the mantis ordinarily stands. The window-corner
is a new and unnatural locale for the insects, but the abundance of
prey here
in summer-time makes it a good feeding-ground.
The family Mantidae includes less than a score of species in this country,
all of them southern in range, and only a few occurring north of the Rio

/*
/-<
130 Cockroaches, Locusts, Grasshoppers, and Crickets

Grande and Gulf coast regions. All the species are carnivorous, and

undoubtedly do much good in making away with many noxious insects. In

1899 some specimens of the common European praying-mantis, Mantis
religiosa (Fig. 161), were found in and
near Rochester, N. Y. They had
probably been accidentally imported
into this country in nursery stocks from
France. As this species seems able
to live farther north than our native

species, Professor Slingerland is laud-

ably trying to establish it in our coun-

try. He takes care of a colony, and
is distributing many of the egg-cases
over the entire country. All the man-
tids lay their eggs in curious masses

(Figs. 162 and 163), covered with a

quickly drying tough mucus. These
egg-cases are attached to branches and
plant-stems in the fall, and the young
hatch in the following summer and
soon grow (moulting several times
and developing wings) to full stature,
which for our most common native
species, Stagmomantis Carolina, is

about z\ inches long.

FIG. 162. Egg-cases of the praying- cl . . ,
.. , ,

mantis, Mantis religiosa. (After Slingerland has collected a num-

Slingerland; natural size.) ber of the old accounts of the Euro-

pean mantis which are of interest as proofs of the light and graceful fancy
of some of the early author-naturalists.
The ancient Greeks gave the insects
the name Mantis, that is, "prophet."
Mouffet, writing over three hundred
years ago, says: "They are called
Mantes, that is, foretellers, either
because by their coming (for they first
of all appear) they do show the
spring
to be at hand, so Anacreon the
poet
sang; or else they foretell death and FIG. 163. Egg-case of praying-mantis,
famine,' as Ca^lius the Scoliast of
Mantis
-T
en>
arrangement of eggs inside.
,S
win
**%*
(Natural
f ^
Theocritus has observed; or, lastly, size.)
because it always holds up its fore feet like hands
praying as it were, after
the manner of their Diviners, who in that gesture did pour out their sup-
 Cockroaches, Locusts, Grasshoppers, and Crickets 131

plications to their Gods." And he says again: "They resemble the Diviners
in the elevation of their hands, so also in likeness of motion; for they do not

sport themselves as others do, nor leap, nor play, but walking softly, they
retain their modesty, and shewes forth a kind of mature gravity. ... So
divine a creature is this esteemed, that if a childe aske the way to such a

place, she will stretch out one of her feet, and shew him the right way, and
seldome or never misse." Piso in his works states that mantids "change into
a green and tender plant, which is of two
hands' breadth. The feet are fixed into
the ground first ;
from these, when neces-

sary, humidity is attracted, roots grow out

and strike the ground;
into thus they
change by degrees, and in a short time
become a perfect plant."
Almost everywhere that mantids occur,
strange superstitions are held concerning
them. Most of these ascribe some degree
of sanctity to them, and to kill them
maliciously is considered sinful. Cowan
says that "the Turks and other Moslems
have been much impressed by the actions
of the common Mantis religiosa, which

greatly resemble some of their own attitudes

of prayer. They readily recognize intelli-
gence and pious intentions in its actions,
and accordingly treat it with respect and
attention, not indeed as in itself an object
of reverence or superstition, but as a fel-
low worshipper of God, whom they believe
that all creatures praise with more or less
consciousness and intelligence. Other su-
perstitions with respect to the Mantis are
current: when it kneelsan angel
it sees
in the way, or hears the rustle of wings its ;

when it alights on your hand you are about

to make the acquaintance of a distin-

guished person; if it alights on your head,

a great honor will shortly be conferred FIG. 164. The walking-stick, Diaphe-
romera jemorata.
upon you. If it
you any way,
injures in
which it does but seldom, you will lose a valued friend by
calumny. Never
kill a Mantis, as it bears charm
against evil." Finally, monkish legends
teli us, says Slingerland, that St. Francis
Xavier, seeing a Mantis moving
132 Cockroaches, Locusts, Grasshoppers, and Crickets

along in its solemn way, holding up itstwo fore legs as in the act of devo-
tion, desired it to sing the praise of God, whereupon the insect carolled
forth a fine canticle!
More amazing than the Mantids for modification of form and appear-
ance away from the usual insect type are the members of the family Phas-
midae. The only representatives of this family in the United States are
the walking-sticks, or twig insects (Fig. 164), of which half a dozen genera,
with from one to three species each, have been recorded. The only one
of these genera which is found in the East is Diapheromera, of which D.

jemorata is the common species. Our other Phasmids are found in the
West or extreme South. All of our species are wingless and are generally
sluggish inmovement, and depend for protection largely on their amazingly
faithfulresemblance in shape and color to twigs, and on their capacity to
emit an ill-smelling fluid from certain glands on their prothorax. Diaphero-
mera femorata (Fig. 164) feeds on the leaves of oaks, walnuts, and probably
other trees. It drops its hundred seed-like eggs loosely and singly on the
ground, where they lie through the winter, hatching irregularly through
the following summer. Some may even go over a second winter before
hatching. Femorata may be either brown or green; so it frequents dead
or leafless, or live and green-leaved parts, according to the correspondence
of its body color with the one or the other of these environments. The long,
slender, wingless body, the thin, long legs held angularly, and the harmonizing
body color, all serve to make the walking-stick well-nigh indistinguishable
when at rest on the twigs.
In tropic and subtropic countries the Phasmids are numerous (over 600
species are known) and present other striking resemblances to the details
of their habitual environment. A conspicuous and perfect example of
resemblance is the green leaf-insect Phyllium (PL XIII, Fig. 2), whose wings,
flattened body,and expanded plate-like legs, head, and prothorax, all bright
green and flecked irregularly with small yellowish spots, like those made
by the attacks of fungi on live leaves, combine to simulate with wonderful
effect a green leaf.

Other examples of such protective resemblance and a discussion of the

origin and significance of the phenomenon may be found in Chapter XVII
of this book.
The genera of Phasmidae occurring in the United States may be distin-
guished by the following key:

Tibiae with a groove at tip to receive the base of the tarsi when bent upon them.
Antennas with less than twenty segments, and much shorter than the fore femora.
BACILLUS.
 Cockroaches, Locusts, Grasshoppers, and Crickets 133
Antennae with many segments, and longer than the fore femora.
Mesothorax twice as long as prothorax ANISOMORPHA.
Mesothorax no longer than prothorax TINEMA.
Tibiae without groove at tip, as above described.
Hind femora with one or more distinct spines on the median line of the under side
near the tip DIAPHEROMERA.
Hind femora without such spines.
Head, especially in female, with a pair of tubercles or ridges on the front between
the eyes SERMYLE.
Head without such tubercle or ridges BACUNCULUS.

One day in early summer of the Centennial Year (1876) the people all
over Kansas might have been seen staring hard with shaded eyes and serious
faces up towards the sun. By persistent looking one could see high in the
air a thin silvery white shifting cloud or haze of which old residents sadly

said, "It's them again, all right." Now this meant, if it were true, that,
far from being was about as wrong as it could be for Kansas.
all right, it

"Them" meant the hateful Rocky Mountain locusts, and the locusts meant
devastation and ruin for Kansas crops and farmers. In 1866 and again
in 1874 and 1875 the locusts had come; first a thin silvery cloud high over-
head sunlight glancing from millions of thin membranous fluttering
wings and then a swarming, crawling, leaping, and ever and always
busily eating horde of locusts over all the green things of the land. And
the old residents spoke the truth in that summer of 1876. It was "them,"

uncounted hosts of them, and only such patriotic farmers as had laid by
money for a rainy day or a grasshopper year could visit the Centennial

Exposition.
Not
all locusts are migratory or appear in such countless swarms as

thisinvader from the high plateau of the northern Rocky Mountains. In

South America another locust species, larger than ours, has similar habits;
having permanent breeding-grounds on the great plateau at the eastern
its

foot of the Chilean Andes and descending almost every year in swarms on
the great wheat-fields of Argentina. And in Algeria and Asia Minor occurs
the migratory locust of the Scriptures, a still other and larger species. But
of the 500 (app.) locust species, members of the family Acridiidae, which
are known in the United States but three or four can be fairly called

migratory, and of these the Rocky Mountain locust, Melanoplus spretus, is

the most conspicuous. The lesser migratory locust, Melanoplus atlanis,
does much injury in New England and other eastern states, while the
pellucid locust, Camnula pellucida, is a migratory species that often does
much harm in California and other western states. Sometimes large
bodies of immature wingless individuals of the large species Dissosteira
longipennis, abundant on the plains of eastern Colorado and western Kansas
 i
34 Cockroaches, Locusts, Grasshoppers, and Crickets

will move slowly on, walking and hopping for

many miles, eating every
green weed and grass-blade in their path, but this is only a limited and
local sort of migration.
Almost all the Acridiidae, despite the many species in the family, are
readily recognizable as locusts
or grasshoppers short-horned
grasshoppers they may be called,
to distinguish them from the
meadow green grasshoppers with
long thread-like antennae because
of their general similarity in ap-
and habit. The body
Locust from lateral aspect (left wings pearance
. ,
FIG. 105. *
f
removed), showing (ao.*) auditory organ, is rather robust, the head is set
(Natural size.) with its long axig at right angles
with the axis of the body, so that the mouth withits strong biting and

crushing jaws is directed downwards (Fig. 165); the antennae are never
as long as the body and are composed of not more than twenty-five

segments; the prothorax is covered laterally as well as dorsally by its large

saddle-like horny pronotum, which projects so as also to cover and protect
from the sharp grass-blades the soft thin-walled neck and the equally
thin- walled suture between prothorax and meso thorax; the abdomen is

broadly and closely joined to the metathorax, and

in the female ends in a short and strong ovipositor
composed of four horny pointed pieces; the hind

legs are much larger than the others and fitted

for leaping, and the fore wings, called tegmina,
are and straight-margined, and serve
narrow
cover and protect the much larger
specially to
thin membranous hind wings, which are plaited
and folded like a fan when the locust is at rest.
The sounds or stridulation of locusts are
made in two ways. When at rest certain species
draw the hind legs up and down across the wing-
covers so that numerous
fine little ridges on the
inner surface broad femora are rasped FlG l66 ._ Locu J
of the
impaled on
.

across a thickened and ridged longitudinal vein thorn by shrike (butcher-

on the outer surface of the wing-covers. When bird) ( Natural size -)
'

in flight certain locusts rub or strike together the upper surface of the
front edge of the hind wings and the under surface of the fore wings
or tegmina. This produces a loud, sharp clacking which can be heard
for a distance of several rods. The loudest "clacking" of this kind
 Cockroaches, Locusts, Grasshoppers, and Crickets 135

that I have heard is made by a species of Trimerotropis, abundant in

the beautiful little glacial "parks" of the Colorado Rockies. Locusts
undoubtedly make sounds to be heard by each other, and it is not difficult
to find in them a matter of more difficulty in most other insects certain

organs which are almost certainly auditory organs, or ears. On the outer
faces of the upper part of the first abdominal segment is a pair of sub-

FIG. 167. The red-legged locust, Melanoplus femur-rubrum, female.

(After Lugger; natural size indicated by line.)

circular clear window-like spots (Figs. 165 and 55). These are thin places
in the body- wall serving as tympana; on the inner face of each is a small

vesicle, and from it a tiny nerve runs to a small auditory ganglion (nerve-

center) at one side of the tympanum. From this auditory ganglion a nerve
runs to the large ventral ganglion in the third thoracic segment. Similar
auditory organs are found in the other singing Orthoptera, the crickets and
katydids, but situated in the front legs instead of on the back.
136 Cockroaches, Locusts, Grasshoppers, and Crickets

The life-history of all our locusts is, in general characteristics, very similar.
The eggs are deposited in oval or bean-shaped packets enclosed in a glutin-
ous substance. They are usually laid just below the surface of the soil,

but in some cases are simply pushed to the ground among the stems of
grasses, while a few locust-species thrust them into soft wood. The strong,
horny ovipositor at the tip of the abdomen is worked into the ground, the
four pieces separated, and the eggs and covering mucous material extruded.
The eggs in a single mass number from twenty-five to one hundred and
twenty-five, varying with different species, and the females of some species
lay several masses. The different species also select different times and
places for egg-laying, some ovipositing in the fall and some in the spring,
while some select hard, gravelly, or sandy spots or well-traveled roads, and
others choose pastures and meadows and the uncultivated margins of irriga-
tion-ditches.
If the eggs are laid in the fall, the more usual case, they do not hatch
until the following spring. The young hoppers are of course wingless, very

small, and pale-colored, but they have the general body make-up of their
parents, with the biting mouth and long-leaping hind legs. They push
their way above ground and feed, as do the adults, on the green foliage of

grasses, herbs, or trees, and in two or three months become full grown and
mature, having moulted five or six times during this growth and developed
wings. The wings begin to appear as minute scale-like projections from
the posterior margins of the back of the meso- and meta-thoracic segments,
and with each moulting are notably larger and more wing-like in appear-
ance. During all this development the wing-pads are so rotated that the
hinder wings (always underneath the fore wings in the adult locust) lie out-
side of and above the fore wings (Fig. 156).
The family Acridiidae includes in the United States about 500 species,
representing 107 genera. These genera are grouped in four subfamilies
as follows:
KEY TO SUBFAMILIES OF ACRIDIID^E.
Pronotum (dorsal wall of prothorax) extending back over the abdomen nearly or quite
to its tip; tegmina (fore wings) short and scale-like .............. TETTIGIN^E.
Pronotum not extending back over abdomen or only slightly; tegmina usually well
developed (sometimes short or wanting).
Prosternum (ventral aspect of prothorax) with a prominent thick conical or cylindrical
spine ......................................................... ACRIDIIN.E.
Prosternum not spined (sometimes a short, oblique, inconspicuous, obtuse tubercle).
Face very oblique .............................................. TRYXALIN^E.
Face nearly or quite vertical

In the subfamily Acridiinae the most conspicuous and economically

important member is the Rocky Mountain or hateful migratory locust,
 Cockroaches, Locusts, Grasshoppers, and Crickets 137

Melanoplus spretus. The invasions of the grain-growing Mississippi Valley

states by this species have been already mentioned. In 1866, 1874, and
1876 such invasions occurred, and before these still others. "Kansas grass-
hoppers" had gained a notoriety which spelled ruin to the state. But,
strangely, these grasshoppers, or locusts, not only were not Kansas born,
but could not even adopt Kansas as a home. The Rocky Mountain locust

FIG. 168. FIG. 169.

FIG. 168. The lesser migratory locust, Melanoplus atlanis, female. (After Lugger;
natural size indicated by line.)
FIG. 169. The differential locust, Melanoplus differentials, female. (After Lugger;
natural size indicated by line.)

has permanent breeding-grounds on the plains and plateaus of Colorado,

its

Idaho, Wyoming, Montana, and British Columbia, at an altitude of from

2000 to 10,000 feet above sea-level, and while able to maintain itself for
a generation or two in the low, moist Mississippi Valley, cannot take up
any permanent residence there. But in those days there were few ranches
and farms on the great plains, and succulent corn and wheat were not at
138 Cockroaches, Locusts, Grasshoppers, and Crickets

hand to feed the millions of young which hatched each spring. So, after
exhausting the scanty wild herbage of their breeding-grounds, and develop-
ing to their winged stage, hosts of locusts would rise high into the air until
they were caught by the great wind-streams bearing southeast, and, with
parachute-like wings expanded and air-sacs in the body stretched to their
fullest, would be borne for a thousand miles to the rich grain-fields of the

FIG. 170. The two-striped locust, Melanoplus bivittatus, female.

(After Lugger; natural size indicated by line.)

Mississippi Valley. As far east as the middle of Iowa and Missouri and
south to Texas these great swarms would spread; and once settled to ground
and started at their chief business, that of eating, not a green thing escaped.
First the grains and grasses; then the vegetables and bushes; then the
leaves and fresh twigs and bark of trees! A steady munching was audible
over the doomed land! And this munching was the devouring of dollars.
Fifty millions of dollars were eaten in the seasons of 1874-76 alone.
 Cockroaches, Locusts, Grasshoppers, and Crickets 139

Remedies there were none; when the summer hosts laid

practically
their eggs in the ground one generation that could be reared in the
for the
invaded land, these eggs could be plowed up, a remedy that is used with
much success in the far western locust-infested states; also when the wingless

young "hoppers" appeared in the spring they could be crushed by heavy

FIG. 171. The American locust, Schistocerca americana, female.

(After Lugger; natural size.)

rollersdrawn across the fields by horses, or burned by scattering straw over

the helpless host and lighting it. Both of these remedies are also used in
western locust-fighting. But against the winged adults there is little that
can be done.
In Asia and South America, where there are also migratory locusts (of
different,much larger species) the natives sometimes try to frighten away
an alighting swarm by smoke and noise, but such a swarm as that which
passed over the Red Sea in November, 1889, spread out for over 2000 square
140 Cockroaches, Locusts, Grasshoppers, and Crickets

miles in area, would be little affected by a bonfire. In Cyprus in 1881,

1300 tons of locust-eggs were destroyed; how many eggs go to make a ton
one can only faintly conceive of.
There has been no serious Rocky Mountain locust invasion of the Missis-
sippi Valley since 1876, and there
will probably never be another. The
locust is being both fed and fought in its own breeding range; many are

FIG. 172. FIG. 175.

FIG. 172. The emarginate locust, Schistocerca emarginata, male. (After Lugger; nat-
ural size.)
FIG. 173. The pale-green locust, Hesperotettix pratensis, female. (After Lugger;
natural size indicated by line.)
FIG; 174. The short-winged locust, Stenobothrus curtipennis, female. (After Lugger;
natural size indicated by line.)
FIG. 175. The sprinkled locust, Chlcealtis conspersa, male. (After Lugger; natural size
indicated by line.)

killed every year, and for those that are left there is food enough and to spare
in the great grain-fields of the northwest plains.
The genus Melanoplus, to which the Rocky Mountain locust belongs,
is the largest of all our Acridiid genera, one hundred and twenty species
found in the United States belonging to it. Of these species a very common
one all over the country is the red-legged locust, Melanoplus femur-rubrum
(Fig. which is about one inch long, with olivaceous brownish body,
167),
clear hind wings and brownish fore wings that have an inconspicuous
longitudinal median series of black spots in the basal half (these spots
 Cockroaches, Locusts, Grasshoppers, and Crickets 141

sometimes wanting). The hind tibiae are normally red (sometimes yellow-
ish), hence the name, although these red hind legs are common to many
other locust species. The lesser migratory locust, M. atlanis (Fig. 168),
is a species same size and appearance which sometimes
of about the

appears in great swarms and does much injury to crops. The largest
species of the genus is M. differentiates (Fig. 169), over an inch and a half long,
with brownish-yellow body, fore wings without spots, and hind wings clear.
It is common in the Southwest, where, in company with M. bivittatus (Fig.

170), nearly as large but readily distinguished by the pair of longitudinal

FIG. 176. FIG. 177. FIG. 178.

FIG. 176. The short-winged green locust, Dichromorpha viridis, female. (After Lugger;
natural size indicated by line.)
FIG. 177. The spotted-winged locust, Orphula pelidina. (After Lugger; natural size
of male 16-19 mm., of female, 20-24 mm.)
FiG. 178. The Carolina locust, Dissosteira Carolina, female. (After Lugger; natural
size indicated by line.)

pale-yellowish stripes extending from the head across the thorax and along the
folded wing-covers nearly to their tips, it often becomes sufficiently abundant
to do serious
injury. These two species are always to be found commonly
in western Kansas, and bivittatus ranges far to the north, being one
of Minnesota's destructive species.

Among the other genera of the subfamily Acridiinae Schistocerca is con-

spicuous because of the large size and wide distribution of its species. The
American locust, S. americana (Fig. 171), measures three inches from head
to tips of tegmina, with reddish-brown body and a longitudinal yellowish
strip extending along the head, thorax, and closed tegmina nearly to their
142 Cockroaches, Locusts, Grasshoppers, and Crickets

tips. The tegmina are opaque and reddish at base, subtransparent dis-
tally; the great hind wings are clear and transparent. This locust is com-
mon in the South, where it sometimes assumes a migratory habit and
becomes very injurious to crops. The leather-colored locust, S. alutaceum,
with dirty brownish-yellow body and paler stripe on top of head and thorax,

FIG. 179. The coral-winged locust, Hippiscus tuberculatus, female. (After Lugger;
natural size indicated by line.)

semi-transparent tegmina, and clear transparent hind wings, and the rusty
locust, S. rubiginosum, with light dust-red body and opaque tegmina, are
the common eastern representatives of this genus. Both are large and
striking forms.
The subfamily Tryxalinae includes a number of locusts distinguished

by the sharp oblique sloping of the face, and in some cases by the much
prolonged and pointed vertex (region of the head between the eyes). In
the East the short-winged locust,
Stenobothrus curtipennis (Fig. 174),
recognizable by its short narrow
wings, yellow under-body, and prom-
inent yellowish hind legs with black

knees, is a common example of this

group. It likes to hide among tall

grasses, where its sprightly tumbling

FIG. 180. Young coral-winged locust, and dodging usually save it from
Hippiscus tuberculatus. (After Lugger; cap ture despite its poor flying and
natural size indicated by line.) , . ,.

leaping powers. The sprinkled

locust, Chloealtis cons per sa (Fig. 175), is an abundant species through-
out the East. It is light reddish brown sprinkled with black spots,

and has* pale yellowish-brown tegmina with many small dark-brown spots,
the wings being clear; it is about three-fourths of an inch long. The
males have the sides of the pronotum shining black. This locust lays its
eggs in rotten stumps or other slightly decayed wood. Blatchley discovered
a female in the act of boring a hole for her eggs in the upper edge of the

topmost board of a six-rail fence. One of the most grotesque of all the
locusts is a member of this subfamily named Achurum brevipenne. The
body is very long and thin, measuring an inch and a half in length by one-
 Cockroaches, Locusts, Grasshoppers, and Crickets 143

tenth of an inch wide in the broadest part; the head is pointed and pro-

jects far forward and upward, the face being very oblique. The wings
are short and the body color brown. Comstock found this locust quite
common on the "wire-grass" which grows in the sand among
in Florida
the saw-palmettoes, and "so closely did their brown linear bodies resemble

dry grass that it was very difficult to perceive them." So the grotesqueness
has its use.
The subfamily CEdipodinas is well represented in the United States,

FIG. 181. Hippiscus tigrinus, female. (After Lugger; nat. size indicated by line.)

containing twenty-four genera and about 140 species. Almost all the familiar
locusts with showy colored hind wings belong to this subfamily. One
of the commonest species all over the United States and Canada is the
Carolina locust, Dissosteira Carolina (Fig. 178), easily recognized by its
black hind wings with broad yellow or yellowish-white margin covered with
dusky spots at the tip. Its body color is pale yellowish or reddish brown,
and it measures 1^-2 inches in length. It flies well; the males have the

habit of hovering in the air a few feet above the ground and making a loud
144 Cockroaches, Locusts, Grasshoppers, and Crickets

"clacking." The species of Hippiscus are heavy, broad-bodied forms

with wings reddish or yellow-
ish at base, then a broad black-
ish band, and the
apex and
margin clear. The fore wings
and body are yellowish to
brown, with darker blotches
and speckles. H. discoideus,
with wings red on basal half,
is common in the East. H.
tuberculatus (Figs. 179 and 180),
the coral-winged locust, or king

grasshopper, also with red

wing-disks, is common in the

Mississippi Valley it makes ;

a very loud rattling while in

FlG. 182. The
yellow- winged locust, Arphia
sulphured. (After Lugger; natural size of
the air. The genus Arphia,
male 23-26 mm., of female 28-30 mm.) also characterized by wings
with bright red
or yellowish
disks but having the fore wings
without large spots or blotches,
usually not even speckled, and
with the body slenderer than
in Hippiscus, comprises about

twenty species scattered over

the whole country. A. xan-
thoptera, with plain smoky
brown fore wings and upper
body, and hind wings with
bright yellow disk, broad smoky
outer band and clearer apex,
is common in the East; A.
tenebrosa (Fig. 183), with brown
and clayey-speckled fore wings
and upper body and hind
wings with coral-red disk and
smoky broad outer band fad-
ing out in apex, is common
FIG. 183. Arphia tenebrosa. (After Lugger; nat- in the West. The green-
ural size indicated by line.).
striped locust, Chortophaga
viridifasciata (Figs. 184 and 185), abundant and familiar in the East and
Mississippi Valley, appears in two forms; in one, the head, thorax, and
 Cockroaches, Locusts, Grasshoppers, and Crickets 145

FIG. 184. The green-striped locust, Chortophaga viridifasciata, form virginiana, female.
(After Lugger; natural size indicated by line.)

FIG. 186. FIG. 187.

FIG. 185. The green-striped locust, Chortophaga viridifasciata, form virginiana, male.
(After Lugger; natural size indicated by line.)
FIG. 186. The clouded locust, Encoptolophus sordidus, male. (After Lugger; nat-
ural size indicated by line.)
FIG. 187. The pellucid locust, Camnula pellucida, female. (After Lugger; natural
size indicated by line.)
146 Cockroaches, Locusts, Grasshoppers, and Crickets

femora are green and there is a broad green stripe on each wing-cover;
the other form is dusky brown all over; both are about i inch (male) to i^
inches (female) long, and have a distinct sharp little median crest on the

FIG. 190. FIG. 191.

FIG. 188. Barren-ground locust, Spharagemon bolli, male. (After Lugger; natural size
male 2022 mm., of female 27-33 mm.)
of
FIG. 189. Spharagemon collare, race scudderi, male. (After Lugger; natural size in-
dicated by line.)
FIG. 190. The long-horned locust, Psinidia jenestralis, male. (After Lugger; natural
size indicated by line.)
FlG. 191. Circolettix verruculatus, male. (After Lugger; natural size indicated by line.)

pronotum. The clouded locust, Encoptolophus sordidus (Fig. i86),is another

species very common in the fall; it is about an inch long, dusky brown

mottled with darker spots; the wing-covers are blotched and the wings
 Cockroaches, Locusts, Grasshoppers, and Crickets 147
clear and transparent; the prothorax looked at from above appears to be

"pinched" at its middle. The males make a loud crackling when in the
air.

It is familiar knowledge that locusts which are readily seen in the air
are extremely difficultto distinguish when alighted. This concealment,
resulting from a harmonizing of the body color with that of the grass or
soil, is of course an advantage to the locust in its "struggle for existence"
and is technically known as protective resemblance (see Chapter XVII). No
locusts show this protective resemblance better
than the species of Trimerotropis (Fig. 193)
especially familiar in the western states. The
colors of various individuals of a single species

vary with the soil colors of the locality, ranging

from whitish to
brownish to slaty
and bluish. I have
taken series of spe-
cimens of Trimero-
tropis sp. in Colorado
showing this whole
range of ground
coloration.

FIG. 192. FIG. 193.

FIG. 192. Mestobregma cincta, male. (After Lugger; natural size indicated by line.)
FIG. 193. The maritime locust, Trimerotropis maritima, female. (After Lugger; nat-
ural size indicated by line.)

The subfamily Tettiginae includes the strange little Acridiids known as

"grouse-locusts." They are all under most of

inch in length, and
them are less than ^ inch. They have the wing-covers reduced to mere
scales, but the pronotum is so long that it extends back over the rest of the
148 Cockroaches, Locusts, Grasshoppers, and Crickets

thorax to the abdomen and more or less covers it. In some species the
pronotum actually extends beyond the tip of the abdomen. The head is

deeply set in the prothorax, the

prosternum being expanded into a broad
border which nearly covers the mouth. As all the grouse-locusts are dark-
colored and without any conspicuous markings, and choose for habitat the
dark ground along streams and ponds, or swampy meadows, they are

FIG. 194. FIG. i

(,5. FIG. 196.
FIG. 194. Nomotettix parvus. (After Lugger; natural size indicated by line.)
FIG. 195. Tettigidea lateralis. (After Lugger; natural size indicated by line.)
FIG. 196. Tettix granulatus, and pronota of two varieties. (After Lugger; natural size
indicated by line.)

They vary much in colora-

infrequently seen except by persistent students.
tion and markings, and harmonize thoroughly with the soil on which
slight

they habitually live. They feed on lichens, moulds, germinating seeds,

and sprouting grasses, and are said to eat
surface mud and muck containing or largely
consisting of decaying vegetable matter. The
eggs are laid in a pear-shaped mass in a
shallow burrow; in May and June the young
hatch in from sixteen to twenty-five days,
becoming mature in late fall, or sometimes
not until the following spring. The nymphs
and adults hibernate, becoming active again
early in spring. A common species is Tettix

granulatus (Fig. 196), slender, length about

FIG. 197. FIG. 198.
FIG. 197. Tettix ornatus. (After inch, and with the narrow pointed pronotum
Lugger; natural size indicated extending beyond the abdomen. This species
7 hibernates among rubbish and loose bark, but
Fie &'.'-Paratettix cucullatus.
(After Lugger; natural size is more or less active on warm winter days.
indicated by line.)
It j s plentiful all through the rest of the year

on its feeding-grounds. T. ornatus (Fig. 197) is a shorter, more robust

species, and is marked with black spots and indefinite yellow blotches as
 Cockroaches, Locusts, Grasshoppers, and Crickets 149

indicated in the figure. In the genus Tettigidea the antennae have from
15 to 22 segments, while in Tettix they have only 12 to 14 segments. Tet-

tigidea lateralis (Fig. 195) is a common species yellowish brown in color,

more yellowish underneath. It is rather robust and the pronotum extends
beyond the tip of the abdomen.
Included in the family Locustidae are katydids, meadow grasshoppers,
cave-crickets, wingless crickets, western crickets, Jerusalem crickets, and
what not, but no locusts. The general reader v of natural history should
always keep clearly in mind the
made by natu-
sharp distinction
between "scientific" and
ralists

"vernacular" names. The ver-

nacular name locust is applied
to insects of the family Acri-
diidae, but not to any of the
members of the family whose
scientific name is Locustidae.
Of the Locustids the best
known representatives are un-
doubtedly the katydids. Anna
Botsford Comstock, the nature-
study teacher of Cornell Uni-
versity, introduces them to her
readers as follows: "The
chances are that he who lies
awake of a midsummer night
must listen, whether he wishes
to do so or not, to an oft-

repeated, rasping song that

says, 'Katy did, Katy did; she

did, she didn't,' over and over
again. There is no use of won-
dering what Katy did or didn't
do, for no mortal will ever
know. If, when the dawn
comes, the listener
eyes
J has ,-.
r IG. 199. Broad-winged katydid, and leaf with
sharp enough to discern one of katydid eggs along edge. (Natural size.)
these singers among the leaves of some neighboring tree, never a note of
explanation will he get. The beautiful, finely veined wings folded close
over the body keep the secret hidden, and the long antennae, looking like
threads of living silk, will wave airily above the droll green eyes as much
as to say, 'Wouldn't you like to know?'"
150 Cockroaches, Locusts, Grasshoppers, and Crickets

The katydids are rather large, almost always green insects that live in
trees and shrubs, where they feed upon the leaves and tender twigs, some-
times doing considerable injury. With almost all the other Locustids,
they will also take animal food if accessible, and some
of the ground-

inhabiting forms undoubtedly depend largely on animal substances for

food. The color and form of the wing-covers and body serve to make them
nearly indistinguishable in the foliage, and as they do
not flock together
in numbers, they are not frequently seen. Their love-calls or songs, how-
ever, make the welkin ring at night from
midsummer until thecoming of frost. Few
katydids sing by day: would bring their
it

enemies, the birds, down on them; but as

twilight approaches, the males begin their
shrilling, which is kept up almost constantly
till
daylight. Like the sound-making Acri-
diids the musical Locustids have a pair of
special auditory organs, or ears, for hearing
these love-songs. These ears are tympanal
organs situated one in the base of each fore
tibia (the Acridiid ears are on the upper
part of the first abdominal segment), and
consist of a thin place in the chitinized

body- wall tympanum), a resonance-

(the
chamber and a special arrangement
inside,
of nerves and ganglia. There are several
genera of these Locustids, corresponding to
the distinctions popularly made under the
vernacular names narrow-winged, round-
winged, angular- winged, oblong leaf-winged,
and broad-winged katydids. The true
katydid is one of the last-named forms,

FIG. 200. Broad-winged katydid, the commonest and most wide-spread species
Cyrtophyllus concavus, male. concavus (Fig. 200).
being Cfytophyllus
(After Harris; natural size.)
It is bright dark-green, and is rarely
distinguished when at rest in the foliage, although familiar to all from its

shrill singing. When specimens of katydids are collected and examined,

concavus may be readily distinguished by the fact that its wings are shorter
than the wing-covers, and these latter are very convex and so curved around
the body that their edges meet above and below. The ovipositor of the
female is short, compressed, slightly curved and pointed. This katydid
ismost in evidence in late summer. People disagree about the melody
and alleged charm of the song. Many cannot distinguish the "katydid"
 Cockroaches, Locusts, Grasshoppers, and Crickets 151

syllables, and Scudder, an experienced student of the Orthoptera, says that

the note, which sounds like xr, has a shocking lack of melody, adding that
the poets who have sung its praises must have heard
it at the distance that lends enchantment. The sounds
are made by
the males exclusively, and result from
the rubbing together of the bases of the wing-covers,
which have the veins and membrane specially modified
for this purpose (see Fig. 201). Concavus lays, in the
autumn, flattened dark slate-colored eggs, about inch
long and one-third as wide, in two rows along a twig,
the eggs overlapping a little. These eggs hatch in the
and the Jyoung, like the adults, feed G
following spring, ;
lus concavus sp.
.

on the foliage of the tree.

The oblong leaf-winged and round-winged katydids belong to the genus
Amblycorypha, and they can be readily recognized by the broad, oblong,
and rounded wing-covers, and the strongly curved ovipositor of the female,
with serrated tip. They are grass-green and have the wings longer than
the wing-covers. The oblong leaf- winged species, A. oblongifolia (Fig. 202),
is 2 inches long to tips of folded
wings, while the round -winged
species, A. rotundijolia, is ij in-
ches or less in length. These
katydids prefer bushes and tall

weeds or even grass-clumps to

tree-tops. Oblongifolia is said by
McNeill to make a "quick shuf-
fling sound which resembles
FIG. 202. The oblong leaf -winged katydid, ' '
or
' '

katy katydid very slight-

Amblycorypha oblongifolia, female. (After
ly," while the song of rotun-
Lugger; natural size.)
dijolia is said by Scudder to be
made both day and night without variation and to consist of two to four
notes, sounding like chic-a-chee, run together and repeated generally once
in about five seconds for an indefinite length of time.

FIG. 203. Angular-winged katydid, Microcentrum laurifolium, male.

(After Riley; natural size.)
The angular-winged katydids, genus Microcentrum, are large, numerous,
152 Cockroaches, Locusts, Grasshoppers, and Crickets

and the most familiarly known of all. The best-known species, M. retinervis,
is over 2 inches long (from head to tip of folded wings) the overlapping ;

dorsal parts of the wing-covers form a conspicuous angle with the lateral

parts, hence the name "angular-winged." The ovipositor of the female is

very short, strongly curved, and with a bluntly pointed, finely serrate tip. The
song of M.
laurifolium (Fig. 203) is said to sound like
repeated from tic

eight to twenty times, at the rate of four a second. The eggs, of which each
female lays from 100 to 150 in the fall, are grayish brown, flat, and long-
oval, about \ inch long by \ inch wide, and are glued in double rows along
twigs or on the edges of leaves (Fig. 199). I have found them on thorns
of the honey-locust, and Howard once received "a batch from a western

correspondent which was found on the edge of a freshly laundried collar

which had lain for some time in a bureau drawer." The rows are side by
side, and the flat eggs overlap each other in their own row. The young
hatch in spring and, slowly growing, moulting, and developing wings, reach
full size and maturity by the middle of the summer.

FIG. 2040. FIG. 2046.

FIG. 2040. The fork -tailed katydid, Scudderia jurcata, female. (After Lugger; nat. size.)
FIG. 2046. The fork-tailed katydid, Scudderia jurcata, male. (After Lugger; nat. size.)

The narrow-winged katydids, belonging to the genus Scudderia (Figs. 204-

206), are smaller than the broader- winged kinds, being not more than i
inches in length to tip of folded wing-covers, and the wing-covers are narrow
and of nearly equal width for their whole length. The ovipositor is broad,

FIG. 205. FIG. 206.

FIG. 205. Scudderia pistillata, female. (After Lugger; natural size.)
FIG. 206. Scudderia pistillata, male. (After Lugger; natural size.)

compressed, and curves sharply upward. These insects frequent shrubbery

and bushes, or coarse grasses and weeds along ravines or ponds; also
marshes, cranberry-bogs, and similar wet places. Their flight is noiseless
 Cockroaches, Locusts, Grasshoppers, and Crickets 153

and zigzag, and when pursued they will take to the lower branches of trees,
especially oaks if near by. The males sing somewhat in daytime as well
as at night, and have different calls for the two times. The females lay
their eggs in the edges of leaves, thrusting them in between the upper and
lower cuticle by means of their flattened and pointed ovipositor.
While almost all katydids are green, a few exceptions are known.
Scudder has found certain pink individuals belonging to a species normally
green. In mountain regions a few species of gray- or granite-colored katy-

FIG. 207. The sword-bearer, Conocephalus ensiger, female. (After Lugger; nat. size.)

dids are known, the color here being quite as protective as the green of the
lowland forms, for these mountain species alight to rest on the granite rocks
of the mountainside. I have found these granite katydids in the Sierra
Nevada of California.

FIG. 208. FIG. 209.

FIG. 208. The sword-bearer, Conocephalus ensiger, male. (After Lugger; nat. size.)
FIG. 209. A common meadow grasshopper, Orchelimum vulgare, female. (After
Lugger; natural size indicated by line.)

The meadow grasshoppers are small, katydid-like Locustids, green and

long- winged, with long, slender hind legs and with the characteristic slender
thread-like antennae longer than the body. These antennae readily distinguish
them from any of the locusts (Acridiidae) which may be found in their com-
pany. The meadow green grasshoppers abound in pastures and meadows,
154 Cockroaches, Locusts, Grasshoppers, and Crickets

and they dislike to take to wing, trusting, when alarmed, to spry leaping or
clever wriggling away and hiding among the lush grasses. Their green
color of course aids very much in protecting them from enemies. They
include three common genera, viz.:
Conocephalus (Figs. 207 and 208), or
cone-headed grasshoppers or sword-
bearers with head produced into a
long, pointed, forward-projecting, cone-
process, slender body, and very
like

FIG. 210. A common meadow lon g> slender, straight or angled, sword-
grasshop-
per, Orchelimum vulgare, male. (After like ovipositor