Transcriber’s notes:
Except for the spelling corrections listed below, the text of this
book has been preserved as in the original, including inconsistent
punctuation, hyphenation and accents.
Lepidotera → Lepidoptera
coccoon → cocoon
subtances → substances
Bütchsli → Bütschli
In this plain-text version, paired underscores denote _italicised
text_, paired asterisks denote *bold text*, a ^ (caret) indicates
^superscripted text and an underscore denotes _subscripted text.
Footnotes have been positioned below the relevant paragraphs and
illustration captions adjacent to the relevant text.
THE COCKROACH
An Introduction to the Study of Insects
STUDIES IN COMPARATIVE ANATOMY--III
THE STRUCTURE AND LIFE-HISTORY
OF
THE COCKROACH
(_PERIPLANETA ORIENTALIS_)
An Introduction to the Study of Insects
BY
L. C. MIALL
PROFESSOR OF BIOLOGY IN THE YORKSHIRE COLLEGE, LEEDS
AND
ALFRED DENNY
LECTURER ON BIOLOGY IN THE FIRTH COLLEGE, SHEFFIELD
LONDON: LOVELL REEVE & CO.
LEEDS: RICHARD JACKSON
1886
STUDIES IN COMPARATIVE ANATOMY.
I.--THE SKULL OF THE CROCODILE. A Manual for Students. By Professor
L. C. MIALL. 8vo, 2_s._ 6_d._
II.--THE ANATOMY OF THE INDIAN ELEPHANT. By Professor L. C. MIALL
and F. GREENWOOD. 8vo, 5_s._
III.--THE COCKROACH: An Introduction to the Study of Insects. By
Professor L. C. MIALL and A. DENNY. 8vo, 7_s._ 6_d._
IV.--MEGALICHTHYS; A Ganoid Fish of the Coal Measures. By Professor
L. C. MIALL (_In preparation_).
MAY BE HAD OF
LOVELL REEVE & CO., LONDON;
RICHARD JACKSON, LEEDS.
PREFACE.
That the thorough study of concrete animal types is a necessary
preliminary to good work in Zoology or Comparative Anatomy will now
be granted by all competent judges. At a time when these subjects,
though much lectured upon, were rarely taught, Döllinger, of Würzburg,
found out the right way. He took young students, often singly, and
made them master such animal types as came to hand, thereby teaching
them how to work for themselves, and fixing in their minds a nucleus
of real knowledge, around which more might crystallise. “What do you
want lectures for? Bring any animal and dissect it here,” said he to
Baer, then a young doctor longing to work at Comparative Anatomy.[1]
It was Döllinger who trained Purkinje, Pander, Baer, and Agassiz,
and such fame cannot be heightened by words of praise. In our own
time and country Döllinger’s methods have been practised by Professor
Huxley, whose descriptive guides, such as the Elementary Biology and
the delightful little book on the Crayfish, now make it easy for
every teacher to work on the same lines. From the description of the
Cockroach in Huxley’s Anatomy of Invertebrated Animals came the impulse
which has encouraged us to treat that type at length. It may easily
turn out that in adding some facts and a great many words to his
account, we have diluted what was valuable for its concentration. But
there are students--those, namely, who intend to give serious attention
to Entomology--who will find our explanations deficient rather than
excessive in detail. It is our belief and hope that naturalists will
some day recoil from their extravagant love of words and names, and
turn to structure, development, life-history, and other aspects of the
animal world which have points of contact with the life of man. We have
written for such as desire to study Insects on this side.
[1] Baer’s account of Döllinger is to be found in the Leben und
Schriften von K. E. von Baer, § 8.
Whoever attempts to tell all that is important about a very common
animal will feel his dependence upon other workers. Much of what is
here printed has been told before. The large number of new figures is,
however, some proof that we have worked for ourselves.
It is a pleasant duty to offer our thanks for friendly help received.
Professor Félix Plateau, of Ghent; Mr. Joseph Nusbaum, of Warsaw;
and Mr. S. H. Scudder, of Cambridge, Massachusetts, have very kindly
consented to treat here of those parts of the subject which they have
specially illustrated by their own labours.[2] Mr. E. T. Newton,
of the Jermyn Street Museum, has lent us the wood blocks used to
illustrate one of his papers on the Brain of the Cockroach. A number
of the figures have been very carefully and faithfully drawn for us
by Miss Beatrice Boyle, a student in the Yorkshire College. We are
much indebted to Dr. Murie, the Librarian of the Linnean Society, for
procuring us access to the extensive literature of Insect Anatomy, and
for answering not a few troublesome questions.
[2] Prof. Plateau’s chief communications will be found on pp. 131 and
159; Mr. Nusbaum has furnished the account of the Development of the
Cockroach, pp. 180 to 195; and Mr. Scudder the Geological History of
the Cockroach, chap. xi.
Five articles on the Cockroach were contributed by us to Science Gossip
in 1884, and some of the figures were then engraved and published.
In issuing a book which has been long in hand, but which can never hope
to be complete, we venture to adopt the words already used by Leydig
concerning his Lehrbuch der Histologie:--“Die eigentlich nie fertig
wird, die man aber für fertig erklären muss, wenn man nach Zeit und
Umständen das Möglichste gethan hat.”
CONTENTS.
CHAP. PAGE
I.--WRITINGS ON INSECT ANATOMY 1
II.--THE ZOOLOGICAL POSITION OF THE COCKROACH 9
III.--THE NATURAL HISTORY OF THE COCKROACH 17
IV.--THE OUTER SKELETON 28
V.--THE MUSCLES; THE FAT-BODY AND CŒLOM 71
VI.--THE NERVOUS SYSTEM AND SENSE ORGANS 86
VII.--THE ALIMENTARY CANAL AND ITS APPENDAGES 113
VIII.--THE ORGANS OF CIRCULATION AND RESPIRATION (including
a section on the Respiratory Movements of Insects,
by Prof. Félix Plateau, of Ghent) 133
IX.--REPRODUCTION 167
X.--DEVELOPMENT (including a section on the Embryonic
Development of the Cockroach, by Joseph Nusbaum,
of Warsaw) 181
XI.--THE COCKROACH OF THE PAST, by S. H. Scudder, of
the U.S. Geological Survey 205
APPENDIX:--
PARASITES OF THE COCKROACH.
SENSE OF SMELL IN INSECTS.
⁂ Where the species is not named, it is to be understood that the
figures are drawn from the Cockroach.
LEEDS:
MCCORQUODALE & CO. LIMITED,
BASINGHALL STREET.
STUDIES IN COMPARATIVE ANATOMY.--No. III.
THE COCKROACH.
CHAPTER I.
WRITINGS ON INSECT ANATOMY.
Marcello Malpighi. 1628–1694.
Jan Swammerdam. 1637–1680.
Pierre Lyonnet. 1707–1789.
Hercule Straus-Dürckheim. 1790–1865.
The lovers of minute anatomy have always been specially attracted to
Insects; and it is not hard to tell why. No other animals, perhaps,
exhibit so complex an organisation condensed into so small a body.
We possess, accordingly, a remarkable succession of memoirs on the
structure of single Insects, beginning with the revival of Anatomy in
the 17th century and extending to our own times. The most memorable
of these Insect-monographs bear the names of Malpighi, Swammerdam,
Lyonnet, and Straus-Dürckheim.
_Malpighi on the Silkworm._
Malpighi’s treatise on the Silkworm (1669) is an almost faultless essay
in a new field. No Insect--hardly, indeed, any animal--had then been
carefully described, and all the methods of work had to be discovered.
“This research,” says Malpighi, “was extremely laborious and tedious”
(it occupied about a year) “on account of its novelty, as well as the
minuteness, fragility, and intricacy of the parts, which required a
special manipulation; so that when I had toiled for many months at this
incessant and fatiguing task, I was plagued next autumn with fevers
and inflammation of the eyes. Nevertheless, such was my delight in
the work, so many unsuspected wonders of nature revealing themselves
to me, that I cannot tell it in words.” We must recall the complete
ignorance of Insect-anatomy which then prevailed, and remember that
now for the first time the dorsal vessel, the tracheal system, the
tubular appendages of the stomach, the reproductive organs, and the
structural changes which accompany transformation were observed, to
give any adequate credit to the writer of this masterly study. Treading
a new path, he walks steadily forward, trusting to his own sure eyes
and cautious judgment. The descriptions are brief and simple, the
figures clear, but not rich in detail. There would now be much to
add to Malpighi’s account, but hardly anything to correct. The only
positive mistakes which meet the eye relate to the number of spiracles
and nervous ganglia--mistakes promptly corrected by Swammerdam. Had the
tract De Bombycibus been the one work of its author, this would have
kept his memory bright, but it hardly adds to the fame of the anatomist
who discovered the cellular structure of the lung, the glandular
structure of the liver and kidney, and the sensory papillæ of the skin,
who first saw the blood-corpuscles stream along a vessel, who studied
very early and very completely the minute structure of plants and the
development of the chick, and whose name is rightfully associated with
the mucous layer of the epidermis, the vascular tufts of the kidney,
and the follicles of the spleen, as well as with the urinary tubules of
Insects.
All that we know of Malpighi commands our respect. Precise and rapid
in his work, keen to discover points of real interest, never losing
himself in details, but knowing when he had done enough, he stands
pre-eminent in the crowd of minute anatomists, who are generally
faithful in a few things, but very unfit to be made rulers over many
things. The last distinct glimpse which we get of him is interesting.
Dr. Tancred Robinson, writing to John Ray, from Geneva, April 18th,
1684, tells how he met Malpighi at Bologna. They talked of the origin
of fossils, and Malpighi could not contain himself about Martin
Lister’s foolish hypothesis that fossils were sports of nature. “Just
as I left Bononia,” he continues, “I had a lamentable spectacle of
Malpighi’s house all in flames, occasioned by the negligence of his
old wife. All his pictures, furniture, books, and manuscripts were
burnt. I saw him in the very heat of the calamity, and methought I
never beheld so much Christian patience and philosophy in any man
before; for he comforted his wife, and condoled nothing but the loss of
his papers, which are more lamented than the Alexandrian Library, or
Bartholine’s Bibliothece, at Copenhagen.”[3]
[3] Correspondence of John Ray, p. 142.
_Swammerdam on the Honey Bee._
Swammerdam’s great posthumous work, the Biblia Naturæ, contains about a
dozen life-histories of Insects worked out in more or less detail. Of
these the May-fly (published during the author’s life-time, in 1675) is
the most famous; that on the Honey Bee the most elaborate. Swammerdam
was ten years younger than Malpighi, and knew Malpighi’s treatise on
the Silkworm--a not inconsiderable advantage. His working-life as a
naturalist comes within the ten years between 1663 and 1673; and this
short space of time was darkened by anxiety about money, as well as by
the religious fanaticism, which in the end completely extinguished his
activity. The vast amount of highly-finished work which he accomplished
in these ten years justifies Boerhaave’s rather rhetorical account
of his industry. Unfortunately, Boerhaave, whom we have to thank
not only for a useful sketch of Swammerdam’s life, but also for the
preservation of most of his writings, was only twelve years old when
the great naturalist died, and his account cannot be taken as personal
testimony. Swammerdam, he tells us, worked with a simple microscope and
several powers. His great skill lay in his dexterous use of scissors.
Sometimes he employed tools so fine as to require whetting under the
microscope. He was famous for inflated and injected preparations. As
to his patience, it is enough to say that he would spend whole days
in clearing a single caterpillar. Boerhaave gives us a picture of
Swammerdam at work which the reader does not soon forget. “His labours
were superhuman. Through the day he observed incessantly, and at night
he described and drew what he had seen. By six o’clock in the morning
in summer he began to find enough light to enable him to trace the
minutiæ of natural objects. He was hard at work till noon, in full
sunlight, and bareheaded, so as not to obstruct the light; and his head
streamed with profuse sweat. His eyes, by reason of the blaze of light
and microscopic toil, became so weakened that he could not observe
minute objects in the afternoon, though the light was not less bright
than in the morning, for his eyes were weary, and could no longer
perceive readily.”
Comparing Swammerdam’s account of the Bee with the useful and amply
illustrated memoir of Girdwoyn (Paris, 1876), it is plain that two
centuries have added little to our knowledge of the structure of this
type. Much has been made out since 1675 concerning the life-history
of Bees, but of what was to be discovered by lens and scalpel,
Swammerdam left little indeed to others. It is needless to dwell
upon the omissions of so early an explorer. Swammerdam proved by
dissection that the queen is the mother of the colony, that the drones
are males, and the working-bees neuters; but he did not find out that
the neuters are only imperfect females. In this instance, as in some
others, Swammerdam’s authority served, long after his death, to delay
acceptance of the truth. It is far from a reproach to him that in the
Honey Bee he lit upon an almost inexhaustible subject. In the 17th
century no one suspected that the sexual economy of any animal could be
so complicated as that which has been demonstrated, step by step, in
the Honey Bee.
_Lyonnet on the Goat Moth._
In Lyonnet’s memoir on the larva of the Goat Moth (Traité Anatomique
de la Chenille qui ronge le bois de Saule, 1760[4]) we must not look
for the originality of Malpighi, nor for the wide range of Swammerdam.
One small thing is attempted, and this is accomplished with unerring
fidelity and skill. There is something of display in the delineation
of the four thousand and forty-one muscles of the Caterpillar, and the
author’s skill as a dissector is far beyond his knowledge of animals,
whether live or dead. The dissections of the head are perhaps the most
extraordinary feat, and will never be surpassed. Modern treatises on
Comparative Anatomy continue to reproduce some of these figures, such
as the general view of the viscera, the structure of the leg, and the
digestive tract. Nearly the whole interest of the volume lies in the
plates, for the text is little more than a voluminous explanation of
the figures.
[4] Copies dated 1762 have a plate representing the microscope and
dissecting instruments used by the author.
It is not without surprise that we find that Lyonnet was an amateur,
who had received no regular training either in anatomy or engraving,
and that he had many pursuits besides the delineation of natural
objects. He was brought up for the Protestant ministry, turned to the
bar, and finally became cipher-secretary and confidential translator
to the United Provinces of Holland. He is said to have been skilled in
eight languages. His first published work in Natural History consisted
of remarks and drawings contributed to Lesser’s Insect Theology (1742).
About the same time, Trembley was prosecuting at the Hague his studies
on the freshwater Polyp, and Lyonnet gave him some friendly help in
the work. Those who care to turn to the preface of Trembley’s famous
treatise (Mémoires pour servir à l’histoire des Polypes d’eau douce,
1744) will see how warmly Lyonnet’s services are acknowledged. He made
all the drawings, and engraved eight of them himself, while Trembley
is careful to note that he was not only a skilful draughtsman, but an
acute and experienced observer. When the work was begun, Lyonnet had
never even seen the operation of engraving a plate. Wandelaar, struck
by the beauty of his drawings, persuaded him to try what he could do
with a burin. His first essay was made upon the figure of a Dragon-fly,
next he engraved three Butterflies, and then, without longer
apprenticeship, he proceeded to engrave the plates still required to
complete the memoir on Hydra.
Lyonnet tells us that the larva of the Goat Moth was not quite his
earliest attempt in Insect Anatomy. He began with the Sheep Tick, but
suspecting that the subject would not be popular, he made a fresh
choice for his first memoir. Enough interest was excited by the Traité
Anatomique to call for the fulfilment of a promise made in the preface
that the description of the pupa and imago should follow. But though
Lyonnet continued for some time to fill his portfolio with drawings
and notes, he never published again. Failing eyesight was one ground
of his retirement from work. What he had been able to finish, together
with a considerable mass of miscellaneous notes, illustrated by
fifty-four plates from his own hand, was published, long after his
death, in the Mémoires du Muséum (XVIII.–XX.).
_Straus-Dürckheim on the Cockchafer._
In beauty and exact fidelity Straus-Dürckheim’s memoir on the
Cockchafer (Considérations Générales sur l’Anatomie Comparée des
Animaux Articulés, auxquelles on a joint l’Anatomie Descriptive du
Melolontha vulgaris, 1828) rivals the work of Lyonnet. Insect Anatomy
was no longer a novel subject in 1828, but Straus-Dürckheim was able
to treat it in a new way. Writing under the immediate influence of
Cuvier, he sought to apply that comparative method, which had proved so
fertile in the hands of the master, to the Articulate sub-kingdom. This
conception was realised as fully as the state of zoology at that time
allowed, and the Considérations Générales count as an important step
towards a complete comparative anatomy of Arthropoda. Straus-Dürckheim
had at command a great mass of anatomical facts, much of which had
been accumulated by his own observations. He systematically compares
Insects with other Articulata, Coleoptera with other Insects, and the
Cockchafer with other Coleoptera. Perhaps no one before him had been
perfectly clear as to the morphological equivalence of the appendages
in all parts of the body of Arthropods, and here he was able to extend
the teaching of Savigny. His limitations are those of his time. If in
certain sections we find his collection of facts to be meagre, and his
generalisations nugatory, we must allow for the progress of the last
sixty years--a progress in which Straus-Dürckheim has his share. It is
the work of science continually to remake its syntheses, and no work
becomes antiquated sooner than morphological generalisation.
It is therefore no reproach to Straus-Dürckheim that his treatise
should now be chiefly valuable, not as “Considérations Générales,”
but as the anatomy of the Cockchafer. Long after his theories and
explanations have ceased to be instructive, when the morphology
and physiology of 1828 have become as obsolete as the Ptolemaic
astronomy, the naturalist will study these exquisite delineations
of Insect-structure with something of the pleasure to be found in
examining for the hundredth time a delicate organism familiar to many
generations of microscopic observers.
The fidelity and love of anatomical detail which characterise
the description of the Cockchafer are not less conspicuous in
Straus-Dürckheim’s Anatomie Descriptive du Chat (1846). Both treatises
have become classical.
We have seen how, in Straus-Dürckheim’s hands, Insect anatomy
became comparative. New studies--histology, embryonic development,
and palæontology--have since arisen to complicate the task of the
descriptive anatomist, and it appears to be no longer possible for
one man to complete the history of any animal of elaborate structure
and ancient pedigree. As a method of research the monograph has had
its day. The path of biological discovery now follows an organ or a
function across all zoological boundaries, and it is in the humbler
office of biological teaching that the monograph finds its proper use.
_Later Insect Anatomists._
It is impossible even to glance at the many anatomists who have
illustrated the structure of Insects by studies, less simple in plan,
but not less profitable to science, than those of the monographers. If
we attempt to select two or three names for express mention, it is with
a conviction that others are left whom the student is bound to hold in
equal honour.
Dufour[5] laboured, not unsuccessfully, to construct a General Anatomy
of Insects, which should combine into one view a crowd of particular
facts. The modern reader will gratefully acknowledge his industry and
the beauty of his drawings, but will now and then complain that his
sagacity does not do justice to his diligence.
[5] Dufour. Rech. anat. et phys. sur les Hémiptères (1833) les
Orthoptères, les Hymenoptères et les Neuroptères (1841), et les
Diptères (1851). Mém. de l’Institut, Tom. IV., VII., XI. Also many
memoirs in Ann. des Sci. Nat.
Newport,[6] a naturalist of greater weight and interest, is memorable
for his skill in minute dissection, for his many curious observations
upon the life-history of Insects (see, for example, his memoir on the
Oil-beetle), and especially for his early appreciation of the value of
embryological study.
[6] Newport. Art. “Insecta,” in Cycl. of Anat. and Phys. (1839),
besides many special memoirs in the Phil. and Linn. Trans.
Leydig[7] was the first to occupy fully the new field of Insect
histology, and point out its resources to the physiologist. In all
his works the student finds beauty and exactness of delineation,
suggestiveness in explanation. Leydig’s contributions to Insect anatomy
and physiology, valuable as they are to the specialist, are not
isolated researches, but form part of a new comparative anatomy, based
upon histology. Incomplete so vast a work must necessarily remain, but
it already extends over considerable sections of the animal kingdom.
[7] Leydig. Vom Bau des Thierischen Körpers (1864), Tafeln zur vergl.
Anatomie (1864), Untersuchungen zur Anat. und Histologie der Thiere
(1883), &c., besides many special memoirs in Müller’s Archiv., Zeits.
f. wiss. Zool., Nova Acta, &c.