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FIELD AND GENERAL ORNITHOLOGY

HANDBOOK

OF

FIELD AND GENERAL

OENITHOLOGY

A MANUAL OF THE STEUCTUEE AND
CLASSIFICATION OF BIEDS

WITH INSTEUCTIONS FOR

COLLECTING AND PEESEEVING SPECIMENS

BY

PEOFESSOE ELLIOTT CODES, M.A, M.D., etc.

VICE-PRESIDENT AMERICAN ORNITHOLOGISTS’ UNION ; FOREIGN MEMBER BRITISH

ornithologists’ union ; corresponding member zoological society of LONDON, ETC.

ILLUSTRATED

Honljon

MACMILLAN AND CO.
1890

All rights reserved

Note. — The Publishers beg to give notice that copies of
this English Edition of Dr. Coues’s Book cannot be introduced
into the United States of America.

WELLCOMF i^'STiTUTE
LIB'^ARY

Coll.

wellVlOmec

Call

No.

CP L.-

PUBLISHEES’ PEEFACE

By arrangement with the American publishers of Professor
Coues’s Key to North American Birds, which has been for
many years the standard text -book of Ornithology, we are
enabled to present a new edition of those portions of the
“ Key” which have not less interest for the English than for
the American public.

The present volume consists of two distinct parts.

Part I., entitled Field Ornithology,” contains the necessary
instructions for the observation and collection of birds in the
field, and for the preparation and preservation of specimens
for scientific study in the cabinet.

Part II., entitled “ General Ornithology,” is a technical
treatise on the classification, the zoological characters, and
the anatomical structure of the class of Birds, in which the
examples cited in illustration of the principles of Ornithology
have for the most part been redrawn by the author from
British instead of American birds.

With the further exception of a few verbal changes, and
slight abridgment in one or two places, made by the author in
revising the proofs, the present “ Handbook ” is a reprint of
the portions of the Key” above specified.

...

CONTENTS

PART I

FIELD ORNITHOLOGY

SECTION PAGE

I. Implements foe Collecting, and their Use . . .3

II. Dogs ......... 14

III. Yarious Suggestions and Directions for Field-Work . 15

IV. Hygiene of Collectorship . . . . .28

Y. Registration and Labelling . . . . .33

YI. Instruments, Materials, and Fixtures for preparing Bird-

skins ........ 38

YII. How TO make a Birdskin . . . . . .42

YIII. Miscellaneous Particulars . . . . .68

IX. Collection of Nests and Eggs . . . . .75

X. Care of a Collection . . . . . .82

PART II

GENERAL ORNITHOLOGY

I. Definition of Birds . . . . . . .91

II. Principles and Practice of Classification . . .99

III. Definitions and Descriptions of the Exterior Parts of Birds 123
lY. An Introduction to the Anatomy of Birds . . . 197

PART I

FIELD OENITHOLOGY

BEING

A MANUAL OF INSTEUCTION FOE
COLLECTING, PEEPAEING, AND PEESEEVING BIEDS

FIELD ORNITHOLOGY

Field ornithology must lead the way to systematic and descriptive
ornithology. The study of birds in the field is an indispensable
prerequisite to their scientific study in the library and the museum.
Directions for observing and collecting birds, for preparing and pre-
serving them as objects of natural history, will greatly help the
student to become a successful ornithologist, if he will faithfully
and intelligently observe these rules. It is believed that the practical
instructions which the author has to give will, if followed out,
enable any one who has the least taste or aptitude for such pursuits
to become proficient in the necessary qualifications of the good
working ornithologist. These instructions are derived from the
writer’s own experience, reaching in time over thirty years, and
extending in area over large portions of North America. Having
made in the field the personal acquaintance of most species of North
American birds, and having shot and skinned with his own hands
several thousand specimens, he may reasonably venture to speak
with confidence, if not also with authority, respecting methods of
study and manipulation. Feeling so much at home in the field —
with his gun for destroying birds, and his instruments for preserv-
ing their skins — he wishes to put the most inexperienced student
equally at ease ; and therefore begs to lay formality aside, that he
may address the reader as if chatting with a friend on a subject of
mutual interest.

§ 1.— IMPLEMENTS FOR COLLECTING, AND THEIR USE

The Double-barrelled Shot Gun is your main reliance. Under
some circumstances you may trap or snare birds, catch them with
bird-lime, or use other devices ; but such cases are exceptions to

4

FIELD ORNITHOLOGY

PART I

the rule that you will shoot birds, and for this purpose no weapon
compares with the one just mentioned. The soul of good advice
respecting the selection of a gun is, Get the test one you can afford to
buy ; go the full length of your purse in the matters of material
and workmanship. To say nothing of the prime requisite, safety,
or of the next most desirable quality, efficiency, the durability of
a high-priced gun makes it cheapest in the end. Style of finish
is obviously of little consequence, except as an index of other
qualities ; for inferior guns rarely, if ever, display the exquisite
appointments that mark a first-rate arm. There is really so little
choice among good guns that nothing need be said on this score ;
you cannot miss it if you pay enough to any reputable maker or
reliable dealer. But collecting is a specialty, and some guns are
better adapted than others to your particular purpose. This is the
destruction, as a rule, of small birds, at moderate range, with the
least possible injury to their plumage. Probably three-fourths or
more of the birds of any miscellaneous collection average under the
size of a pigeon, and were shot within thirty yards. A heavy gun
is therefore unnecessary, in fact ineligible, the extra weight being
useless. You will find a gun of seven and a half to eight pounds
weight most suitable. For similar reasons the bore should be
small ; I prefer fourteen gauge, and should not think of going over
twelve. Length of barrel is of less consequence than many suppose ;
for myself, I incline to a rather long barrel — one nearer thirty-three
than twenty-eight inches — believing that such a barrel may throw
shot better ; but I am not sure that this is even the rule, while
it is well known that several circumstances of loading, besides some
almost inappreciable differences in the way barrels are bored, will
cause guns apparently exactly alike to throw shot differently.
Length and crook of stock should of course be adapted to your
figure — a gun may be made to fit you, as well as a coat. For wild-
fowl shooting, and on some other special occasions, a heavier and
altogether more powerful gun will be preferable.

Breech-loader v. Muzzle-loader, a case formerly argued, has
long been settled in favour of the former. Provided the mechanism
and workmanship of the breech be what they should, there are no
valid objections to offset obvious advantages, some of which are
these ; ease and rapidity of loading, and consequent delivery of
shots in quick succession; facility of cleaning; compactness and
portability of ammunition ; readiness with which different- sized
shot may be used. This last is highly important to the collector,
who never knows the moment he may wish to fire at a very
different bird from such as he has already loaded for. The muzzle-
loader must always contain the fine shot with which nine-tenths
of your specimens will be secured ; if in both barrels, you cannot

SEC. I IMPLEMENTS FOR COLLECTING, AND THEIR USE 5

deal with a hawk or other large bird with reasonable prospects of
success j if in only one barrel, the other being more heavily charged,
you are crippled to the extent of exactly one-half of your resources
for ordinary shooting. Whereas, with the breech-loader you will
habitually use mustard-seed in both barrels, and yet can slip in a
different shell in time to seize most opportunities requiring large
shot. This consideration alone should decide the case. Moreover,
the time spent in the field in loading an ordinary gun is no small
item ; while cartridges may be charged in your leisure at home.
This should become the natural occupation of your spare moments.
No time is really gained ; you simply change to advantage the time
consumed. Metal shells, charged with loose ammunition, and
susceptible of being reloaded many times, may be used instead of
any special fixed ammunition which, once exhausted in a distant
place (and circumstances may upset the best calculations on that
score), leaves the gun useless. On charging the shells, mark the
number of the shot used on the outside wad ; or better, use
coloured wads — say plain white for dust shot, and red, blue, and
green for certain other sizes. If going far away; take as many
shells as you think can possibly be wanted — and a few more.

Experience, however, will soon teach you to prefer paper
cartridges for breech-loaders. They may of course be loaded
according to circumstances, with the same facility as metal shells, and
even reloaded if desired. It is a good deal of trouble to take care
of metal shells, to prevent loss, keep them clean, and avoid bending
or indenting ; while there is often a practical difficulty in recapping
— at least with the common styles that take a special primer.
Those fitted with a screw top holding a nipple for ordinary caps are
expensive. Paper cartridges come already capped, so that this
bother is avoided, and it is not ordinarily worth while to reload
them. They are made of different colours, distinguishing various
sizes of shot used without employing the coloured wads otherwise
required. They may be taken into the field empty and loaded on
occasion to suit ; but it is better to pay a trifle extra to have them
loaded at the shop. In such case, about four-fifths of the stock
should contain mustard-seed, nearly all the rest about No. 7, a very
few being reserved for about No. 4. Cost of ammunition is
hardly appreciably increased; its weight is put in the most
conveniently portable shape ; the whole apparatus for carrying it
loose and for loading the shells is dispensed with ; much time is
saved, the entire drudgery (excepting gun- cleaning) of collecting
being avoided. I was prepared in this way during the summer of
1873 for the heaviest work I ever succeeded in accomplishing dur-
ing the same length of time. In June, when birds were plentiful,

I easily averaged fifteen skins a day, and occasionally made twice

6

FIELD ORNITHOLOGY

PART I

as many. As items serving to base calculations, I may mention
that in four months I used about two thousand cartridges, loaded,
at $42 per M., with seven-eighths of an ounce of shot and two and
three -fourths drachms of powder. Only about three hundred
were charged with shot larger than mustard-seed. In estimating
the size of a collection that may result from use of a given number
of cartridges, it may not be safe for even a good shot to count on
much more than half as many specimens as cartridges. The number
is practically reduced by the following steps : Cartridges lost or
damaged, or originally defective ; shots missed ; birds killed or
wounded, not recovered ; specimens secured unfit for preservation,
or not preserved for any reason ; specimens accidentally spoilt in
stuffing, or subsequently damaged, so as to be not worth keeping ;
and finally, use of cartridges to kill game for the table.

Other Weapons, etc. — An ordinary single-larrel gun will of
course answer; but is a sorry makeshift, for it is sometimes so
poorly constructed as to be unsafe, and can at best be only just
half as effective. This remark does not apply to any of the fine
single-barrelled breech-loaders now made. You will find these very
effective weapons, and they are not at all expensive. An arm now
much used by collectors is a kind of breech-loading pistol, with or
without a skeleton gun-stock to screw into the handle, and taking a
particular style of metal cartridge, charged with a few grains of
powder, or with nothing but the fulminate. They are very light,
very cheap, safe and easy to work, and astonishingly effective up to
twenty or thirty yards ; making probably the best “second choice”
after the matchless double-barrelled breech-loader itself. The cane-
gun should be mentioned in this connection. It is a single-barrel,
lacquered to look like a stick, with a brass stopper at the muzzle to
imitate a ferule, countersunk hammer and trigger, and either a
simple curved handle, or a light gunstock-shaped piece that screws
in. The affair is easily mistaken for a cane. Some have acquired
considerable dexterity in its use ; my own experience with it is very
limited and unsatisfactory ; the handle always hit me in the face,
and I generally missed my bird. It has only two recommendations.
If you approve of shooting on Sunday, and yet scruple to shock
popular prejudice, you can slip out of town unsuspected. If you
are shooting where the law forbids destruction of small birds, — a
wise and good law that you may sometimes be inclined to defy, —
artfully careless handling of the deceitful implement may prevent
arrest and fine. A Uow-gun is sometimes used. It is a long slender
tube of wood, metal, or glass, through which clay balls, tiny arrows,
etc., are projected by force of the breath. It must be quite an art
to use such a weapon successfully, and its employment is necessarily
exceptional. Some uncivilised tribes are said to possess marvellous

SEC. I IMPLEMENTS FOR COLLECTING, AND THEIR USE

7

skill in the use of long bamboo blow-guns ; and such people are
often valuable employes of the collector. I have had no experience
with the noiseless air-gun, which is, in effect, a modified blow-gun,
compressed air being the explosive power. Nor can I say much of
various methods of trapping birds that may be practised. On these
points I must leave you to your own devices, with the remark that
horse-hair snares, set over a nest, are often of great service in securing
the parent of eggs that might otherwise remain unidentified. I have
no practical knowledge of bird -lime. A method of netting birds

alive, which I have tried, is both easy and successful. A net of
fine green silk, some eight or ten feet square, is stretched perpen-
dicularly across a narrow part of one of the little brooks, overgrown
with briers and shrubbery, that intersect many of our meadows.
Eetreating to a distance, the collector beats along the shrubbery
making all the noise he can, urging on the little birds till they reach
the almost invisible net and become entangled in trying to fly
through. I have in this manner taken a dozen sparrows and the
like at one “drive.” But the gun can rarely be laid aside for this
or any other device.

Ammunition. — The best powder is that combining strength and
cleanliness in the highest compatible degree. In some brands too
much of the latter is sacrificed to the former. Other things being
equal, a rather coarse powder is preferable, since its slower action
tends to throw shot closer. Some numbers are said to be “too
quick ” for fine breech-loaders. Inexperienced sportsmen and col-
lectors almost invariably use too coarse shot. Then two evils result ;
The number of pellets in a load is decreased, the chances of killing
being correspondingly lessened ; and the plumage is badly injured,
either by direct mutilation, or by subsequent bleeding through large
holes. As already hinted, shot cannot be too fine for your routine
collecting. Use “ mustard-seed,” or “ dust-shot,” as it is variously
called j it is smaller than any of the sizes usually numbered. As
the very finest can only be procured in cities, provide yourself
liberally on leaving any centre of civilisation for even a country
village, to say nothing of remote regions. A small bird that would
have been torn to pieces by a few large pellets, may be riddled with
mustard-seed and yet be preservable ; moreover, there is, as a rule,
little or no bleeding from such minute holes, which close up by the
elasticity of the tissues involved. It is astonishing what large birds
may be brought down with these tiny pellets. I have killed hawks
with such shot, knocked over a wood-ibis at forty yards, and once
shot a wolf dead with No. 10 — though I am bound to say the
animal was within a few feet of me. After dust-shot, and the nearest
number or two, No. 8 or 7 will be found most useful. Water-fowl,
thick-skinned sea-birds like loons, cormorants, and pelicans, and a

FIELD ORNITHOLOGY

PART I

few of the largest land-birds, require heavier shot. I have had no
experience with the substitution of fine gravel or sand, much less
water, as a projectile ; besides shot I never fired anything at a bird
except my ramrod, on one or two occasions, when I never afterward
saw either the bird or the stick. Cut felt wads are the only suitable
article. Ely’s “ chemically prepared ” wadding is the best. It is
well, when using plain wads, occasionally to drive a greased one
through the barrel. Since you may sometimes run out of wads
through an unexpected contingency, always keep a wad-cutter to fit
your gun. You can make serviceable wads of pasteboard, but they
are inferior to felt. Cut them on the flat sawn end of a stick of
firewood. Use a wooden mallet, instead of a hammer or hatchet,
and so save your cutter. Soft paper is next best after wads ; I
have never used rags, cotton, or tow, fearing these tinder-like sub-
stances might leave a spark in the barrels. Crumbled leaves or
grass will answer at a pinch.

Other Equipments. — (a) For the gun. A gun-case will come
cheap in the end, especially if you travel much. The usual box,
divided into compartments, and well lined, is the best, though the
full-length leather or india-rubber cloth case answers very well.
The box should contain a small kit of tools, such as mainspring-vice,
nipple-wrench, screw-driver, etc. A stout hard-wood cleaning-rod,
with wormer, will be required. It is always safe to have parts of
the gun-lock, especially mainspring, in duplicate. For muzzle-
loaders extra nipples and extra ramrod heads and tips often come
into use. For breech-loaders the apparatus for charging the shells
is practically indispensable, (h) For ammunition. Metal shells or
paper cartridges may be carried loose in the large lower coat pocket,
or in a leather satchel. There is said to be a chance of explosion
by some unlucky blow, when they are so carried, but I never knew
of an instance. Another way is to fix them separately in a row in
snug loops of soft leather sewn continuously along a stout waist-belt ;
or in several such horizontal rows on a square piece of thick leather,
to be slung by a strap over the shoulder. But better than anything
else is a stout linen vest, similarly furnished with loops holding each
a cartridge ; this distributes the weight so perfectly, that the usual
“ forty rounds ” may be carried without feeling it. The appliances
for loose ammunition are almost endlessly varied, so every one may
consult his taste or convenience. But now that everybody uses the
breech-loader, shot-pouches and powder-flasks are among the things
that were, (c) For specimens. You must always carry paper in
which to wrap up your specimens, as more particularly directed
beyond. Nothing is better for this purpose than writing-paper ;
“ rejected ” or otherwise useless MSS. may thus be utilised. The
ordinary game-bag, with leather back and network front, answers

SEC. I IMPLEMENTS FOR COLLECTING, AND THEIR USE

9

very well ; but a light basket, fitting the body, such as the creel
used by anglers, is the best thing to carry specimens in. Avoid
putting specimens into pockets, unless you have your coat-tail largely
excavated ; crowding them into a close pocket, where they press
each other, and receive warmth from the person, will injure them.
It is always well to take a little raw cotton into the field, to plug
up shot -holes, mouth, nostrils, or vent, immediately if required.
{d) For yourself. The indications to be fulfilled in your clothing
are these : Adaptability to the weather ; and since a shooting-coat
is not conveniently changed, while an overcoat is ordinarily ineli-
gible, the requirement is best met by different underclothes. Easy
fit, allowing perfect freedom of muscular action, especially of the
arms. Strength of fabric, to resist briers and stand wear ; velveteen
and corduroy are excellent materials. Subdued colour, to render
you as inconspicuous as possible, and to show dirt the least. Multi-
plicity of pockets — a perfect shooting-coat is an ingenious system of
hanging pouches about the person. Broad-soled, low-heeled boots
or shoes, giving a firm tread even when wet. Close-fitting cap with
prominent visor, or low soft felt hat, rather broad-brimmed. Let
india-rubber goods alone ; the field is no place for a sweat-bath.

Qualifications for Success. — With the outfit just indicated
you command all the required appliances that you can huy, and the
rest lies with yourself. Success hangs upon your own exertions ;
upon your energy, industry, and perseverance ; your knowledge
and skill ; your zeal and enthusiasm, in collecting birds, much as
in other affairs of life. But that your efforts — maiden attempts
they must once have been if they be not such now — may be directed
to best advantage, further instructions may not be unacceptable.

To Carry a Gun without peril to human life or limb is the
ah c oi its use. “There’s death in the pot.” Such constant care
is required to avoid accidents that no man can give it by continual
voluntary or conscious effort : safe carriage of the gun must become
an unconscious habit, fixed as the movements of an automaton.
The golden rule and whole secret is : the muzzle must never sweep the
horizon ; accidental discharge should send the shot into the ground
before your feet, or away up in the air. There are several safe and
easy ways of holding a piece ; they will be employed by turns to
relieve particular muscles when fatigued. 1. Hold it in the hollow
of the arm (preferably the left, as you can recover to aim in less
time than from the right), across the front of your person, the hand
on the grip, the muzzle elevated about 45°. 2. Hang it by the

trigger-guard hitched over the forearm brought round to the breast,
the stock passing behind the upper arm, the muzzle pointing to the
ground a pace or so in front of you. 3. Shoulder it, the hand on
the grip or heel-plate, the muzzle pointing upward at least 45°. 4.

10

FIELD ORNITHOLOGY

PART I

Shoulder it reversed, the hand grasping the barrels about their
middle, the muzzle pointing forward and downward ; this is per-
fectly admissible, but is the most awkward position of all to recover
from. Always carry a loaded gun at half-cock^ unless you are about
to shoot. The best guns are now fitted with rebounding locks,
having a device by which the hammer is thrown back to half-cock
as soon as the blow is delivered on the firing-pin. This admirable
device is a great safeguard, and is particularly eligible for breech-
loaders, as the barrels may be unlocked and relocked without
touching the hammers. Unless the lock fail, accidental discharge
is impossible, except under these circumstances : {a) a direct blow
on the nipple or pin ; (h) catching of both hammer and trigger
simultaneously, drawing back of the former and its release whilst
the trigger is still held, — the chances against which are simply
incalculable. Full-cock, ticklish as it seems, is safer than no-cock,
when a tap on the hammer, or a slight catch and release of the
hammer, may cause discharge. Never let the muzzle of a loaded
gun point toward your own person for a single instant. Get your
gun over fences, or into boats or carriages, before you get over or
in yourself, or at any rate no later. Kemove caps or cartridges on
entering a house. Never aim a gun, loaded or not, at any object,
unless you mean to press the trigger. Never put a loaded gun
away long enough to forget whether it is loaded or not. Never
leave a loaded gun to be found by others under circumstances
reasonably presupposing it to be unloaded. Never put a gun where
it can be knocked down by a dog or a child. Never imagine that
there can be any excuse for putting away a breech-loader loaded
under any circumstances. Never forget that the idiots who kill
people because they “didn’t know it was loaded,” are perennial.
Never forget that though a gunning accident may be sometimes
interpreted (from a false standpoint) as a “dispensation of Provi-
dence,” such dispensations happen oftenest to the careless.

To Clean a Gun properly requires some knowledge, more
good temper, and most “ elbow-grease ” ; it is dirty, disagreeable,
inevitable work, which laziness, business, tiredness, indifference, and
good taste will by turns tempt you to shirk. After a hunt you are
tired, have your clothes to change, a meal to eat, a lot of birds to
skin, a journal to write up. If you “sub-let” the contract, the
chances are it is but half fulfilled j serve yourself, if you want to
be well served. If you cannot find time for a regular cleaning, an
intolerably foul gun may be made to do another day’s work by
swabbing for a few moments with a wet (not dripping) rag, and
then with an oiled one. For the full wash use cold water first ] it
loosens dirt better than hot water. Set the barrels in a pail of
water ; wrap the end of the cleaning rod with tow or cloth, and

SEC. I IMPLEMENTS FOR COLLECTING, AND THEIR USE

1 1

pump away till your arms ache. Change the rag or tow, and the
water too, till they both stay clean for all the swabbing you can do.
Then use boiling water till the barrels are well heated ; wipe as
dry as possible inside and out, and set them by a fire. Finish with
a light oiling, inside and out; touch up all the metal about the
stock, and polish the wood-work. Do not remove the locks oftener
than is necessary ; every time they are taken out, something of the
exquisite fitting that marks a good gun may be lost ; as long as
they work smoothly take it for granted they are all right. To
keep a gun well, under long disuse, it should have had a particu-
larly thorough cleaning; the chambers should be packed with
greasy tow ; greased wads may be rammed at intervals along the
barrels ; or the barrels may be filled with melted tallow. Neat’s-
foot is recommended as the best easily procured oil ; the porpoise-
oil which is used by watchmakers is the very best ; the oil made
for use on sewing-machines is excellent ; “ olive ” oil (made of lard)
for table use answers the purpose. The quality of an oil may be
improved by putting in it a few tacks, or scraps of zinc, — the oil
expends its rusting capacity in oxidising the metal. Inferior oils
get “sticky.” One of the best preventives of rust is mercurial
(“ blue ”) ointment ; it may be freely used. Kerosene will remove
rust ; but use it sparingly, for it “ eats ” sound metal too.

To Load a Gun effectively requires something more than
knowledge of the facts that the powder should go in before the
shot, and that each should have a wad atop. The most nearly
universal fault is use of too much shot for the amount of powder ;
and the next, too much of both. The rule is hulk for hulk of powder
and shot. If not exactly this, then rather less shot than powder.
It is absurd to suppose, as some persons do who ought to know
better, that the more shot in a gun the greater the chances of kill-
ing. The projectile force of a charge cannot possibly be greater
than the vis inertim of the gun as held by the shooter. The explo-
sion is manifested in all directions, and blows the shot in one way
simply because it has no other escape. If the resistance in front
of the powder were greater than elsewhere, the shot would not
budge, but the gun would fly backward, or burst. This always
reminds me of Lord Dundreary’s famous conundrum — Why does a
dog wag his tail ? Because he is bigger than his tail ; otherwise
the tail would wag him. A gun shoots shot because the gun is the
heavier ; otherwise the shot would shoot the gun. Every unneces-
sary pellet is a pellet against you, not against the game. The
experienced sportsman uses about one-third less shot than the tyro,
with proportionally better result, other things being equal. As to
powder, moreover, a gun can only burn just so much, and every
grain blown out unburnt is wasted. No express directions for

12

FIELD ORNITHOLOGY

PART I

absolute weight or measures of either powder or shot can be given ;
in fact, different guns take as their most effective charge such a
variable amount of ammunition, that one of the first things you
have to learn about your own arm is, its normal charge -gauge.
Find out, by assiduous target practice, what absolute amounts (and
to a slight degree, what relative proportion) of powder and shot are
required to shoot the farthest and distribute the pellets most evenly.
This practice, furthermore, will acquaint you with your gun’s capa-
cities in every respect. You should learn exactly what it will and
what it will not do, so as to feel perfect confidence in your arm
within a certain range, and to waste no shots in attempting miracles.
Immoderate recoil is a pretty sure sign that the gun is overloaded,
or otherwise wrongly charged ; and all force of recoil is subtracted
from the impulse of the shot. It is useless to ram powder very
hard ; two or three smart taps of the rod will suffice, and more will
not increase the explosive force. On the shot the wad should
simply be pressed close enough to fix the pellets immovably.
These directions apply to the charging of metal or paper cartridges
as well as to loading by the muzzle. The latter operation is rarely
required, now that guns of every grade are made to break at the
breech. Finally, let me impress upon you the expediency of light
loading in your routine collecting. Three-fourths of your shots
need not bring into action the gun’s full powers of execution.
You will shoot more birds under than over .thirty yards ; not a
few you must secure, if at all, at ten or fifteen yards ; and your
object is always to kill them with the least possible damage to the
plumage. I have, on particular occasions, loaded even down to
one third oz. of shot and one and a half dr. of powder. There is
astonishing force compressed in a few grains of powder ; an aston-
ishing number of pellets in the smallest load of mustard-seed.

To Shoot successfully is an art which may be acquired by
practice, and can be learned only in the school of experience. No
general directions will make you a good shot, any more than a
proficient in music or painting. To tell you that in order to hit a
bird you must point the gun at it and press the trigger, is like
saying that to play on the fiddle you must shove the bow across
the strings with one hand while you finger them with the other ;
in either case the result is the same, a noise, but neither music nor
game. Nor is it possible for every one to become an artist in
gunnery; a “crack-shot,” like a poet, is born, not made. For
myself, I make no pretensions to genius in that direction ; for
although I generally make fair bags, and have destroyed many
thousand birds in my time, this is rather owing to some familiarity
I have gained with the habits of birds, and a certain knack, acquired
by long practice, of picking them out of trees and bushes, than to

SEC. I IMPLEMENTS FOR COLLECTING, AND THELR USE 13

skilful shooting from the sportsman’s standpoint ; in fact, if I cut
down two or three birds on the wing without a miss I am working
quite up to my average in that line. But any one not purblind or
a “ butter-fingers ” can become a reasonably fair shot by practice,
and do good collecting. It is not so hard, after all, to sight a gun
correctly on an immovable object, and collecting differs from sport-
ing proper in this, that comparatively few birds are shot on the
wing. But I do not mean to imply that it requires less skill to
collect successfully than to secure game ; on the contrary, it is finer
shooting, I think, to drop a warbler skipping about a tree-top than
to stop a quail at full speed ; while hitting a sparrow that springs
from the grass at one’s feet to dicker in sight a few seconds and
disappear is the most difficult of all shooting. Besides, a crack shot,
as understood, aims unconsciously, with mechanical accuracy and
certitude of hitting ; he simply wills, and the trained muscles obey
without his superintendence, just as the fingers form letters with
the pen in writing ; whereas the collector must usually supervise
his muscles all through the act and see that they mind. In spite
of the proportion of snap-shots of all sorts you will have to take,
your collecting shots, as a rule, are made with deliberate aim.
There is much the same difference, on the whole, between the
sportsman’s work and the collector’s, that there is between shot-gun
and rifle practice, collecting being comparable to the latter. It is
generally understood that the acme of skill with the two weapons
is an incompatibility ; and, certainly, the best shot is not always the
best collector, even supposing the two to be on a par in their know-
ledge of birds’ haunts and habits. Still, a hopelessly poor shot can
only attain fair results by extraordinary diligence and perseverance.
Certain principles of shooting may perhaps be reduced to words.
Aim deliberately directly at an immovable object at fair range.
Hold over a motionless object when far off, as the trajectory of the
shot curves downward. Hold a little to one side of a stationary
object when very near, preferring rather to take the chances of
missing it with the peripheral pellets, than of hopelessly mutilating
it with the main body of the charge. Fire at the first fair aim,
without trying to improve what is good enough already. Never
“ pull ” the trigger, but fress it. Bear the shock of discharge with-
out flinching. In shooting on the wing, fire the instant the heel
of the gun taps your shoulder j you will miss at first, but by and
by the birds will begin to drop, and you will have laid the founda-
tion of good shooting, the knack of “ covering ” a bird unconsciously.
The habit of “ poking ” after a bird on the wing is an almost incur-
able vice, and may keep you a poor shot all your life. (The col-
lector’s frequent necessity of poking after little birds in the bush is
what so often hinders him from acquiring brilliant execution.) Aim

H

FIELD ORNITHOLOGY

PART 1

ahead of a flying bird — the calculation to be made varies, according
to the distance of the object, its velocity, its course and the wind,
from a few inches to several feet ; practice will finally render it
intuitive.

§ 2.— DOGS

A Good Dog is one of the most faithful, respectful, affec-
tionate, and sensible of brutes ; deference to such rare qualities
demands a chapter, however brief. A trained dog is the indis-
pensable servant of the sportsman in his pursuit of most kinds of
game ; but I trust I am guilty of no discourtesy to the noble
animal, when I say that he is a luxury rather than a necessity to
the collector — a pleasant companion, who knows almost everything
except how to talk, who converses with his eyes and ears and tail,
shares comforts and discomforts with equal alacrity, and occasion-
ally makes himself useful. So far as a collector’s work tallies
with that of a sportsman, the dog is equally useful to both ; but
finding and telling of game aside, your dog’s services are restricted
to companionship and retrieving. He may, indeed, flush many
sorts of birds for you ; but he does it, if at all, at random, while
capering about ; for the brute intellect is limited after all, and can-
not comprehend a naturalist. The best trained setter or pointer
that ever marked a quail could not be made to understand what
you are about, and it would ruin him for sporting purposes if he
did. Take a well-bred dog out with you, and the chances are he
will soon trot home in disgust at your performances with jack-
sparrows and tomtits. It implies such a perversion of a good dog’s
instincts to make him really a useful servant of the collector, that I
am half inclined to say nothing about retrieving, and tell you to
make a companion of your dog, or let him alone. I was followed
for several years by “ the best dog I ever saw ” (every one’s gun,
dog, and child is the best ever seen), and a first-rate retriever ; yet
I always preferred, when practicable, to pick up my own birds,
rather than let a delicate plumage into a dog’s mouth, and scolded
away the poor brute so often, that she very properly returned the
compliment, in the end, by retrieving just when she felt like it.
However, we remained the best of friends. Any good setter,
pointer, or spaniel, and some kinds of curs, may be trained to
retrieve. The great point is to teach them not to “ mouth ” a bird ;
it may be accomplished by sticking pins in the ball with which
their early lessons are taught. Such dogs are particularly useful in
bringing birds out of the water, and in searching for them when
lost. One point in training should never be neglected : teach a dog

SEC. Ill SUGGESTIONS AND DIRECTIONS FOR FIELD-WORK 15

what “to heel” means, and make him obey this command. A
riotous brute is simply unendurable under any circumstances.

§ 3.— VARIOUS SUGGESTIONS AND DIRECTIONS FOR
FIELD-WORK

To be a good Colleetop, and nothing more, is a small
affair ; great skill may be acquired in the art, without a single
quality commanding respect. One of the most vulgar, brutal, and
ignorant men I ever knew was a sharp collector and an excellent
taxidermist. Collecting stands much in the same relation to orni-
thology that the useful and indispensable office of an apothecary
bears to the duties of a physician. A field-naturalist is always
more or less of a collector ; the latter is sometimes found to know
almost nothing of natural history worth knowing. The true orni-
thologist goes out to study birds alive and destroys some of them
simply because that is the only way of learning their structure and
technical characters. There is much more about a bird than can
be discovered in its dead body, — how much more, then, than can be
found out from its stuffed skin ! In my humble opinion the man
who only gathers birds, as a miser money, to swell his cabinet, and
that other man who gloats, as miser-like, over the same hoard, both
work on a plane far beneath where the enlightened naturalist stands.
One looks at Nature, and never knows that she is beautiful \ the
other knows she is beautiful, as even a corpse may be ; the
naturalist catches her sentient expression, and knows how beautiful
she is ! I would have you to know and love her ; for fairer mis-
tress never swayed the heart of man. Aim high ! — press on, and
leave the half-way house of mere collectorship far behind in your
pursuit of a delightful study, nor fancy the closet its goal.

Birds may be sought anywhere, at any time ; they should
be sought everywhere, at all times. Some come about your door-
step to tell their stories unasked. Others spring up before you as
you stroll in the field, like the flowers that enticed the feet of
Proserpine. Birds flit by as you measure the tired roadside, lend-
ing a tithe of their life to quicken your dusty steps. They disport
overhead at hide-and-seek with the foliage as you loiter in the shade
of the forest, and their music now answers the sigh of the tree-tops,
now ripples an echo to the voice of the brook. But you will not
always so pluck a thornless rose. Birds hedge themselves about
with a bristling girdle of brier and bramble you cannot break ;
they build their tiny castles in the air surrounded by impassable
moats, and the drawbridges are never down. They crown the

i6

FIELD ORNITHOLOGY

PART I

mountain-top you may lose your breath to climb \ they sprinkle the
desert where your parched lips may find no cooling draught ; they
fleck the snow-wreath when the nipping blast may make you turn
your back ; they breathe unharmed the pestilent vapours of the
swamp that mean disease, if not death, for you ; they outride the
storm at sea that sends strong men to their last account. Where
now will you look for birds %

And yet, as skilled labour is always most productive, so expert
search yields more than random or blundering pursuit. The more
varied the face of a country, the more various its birds. A place
all plain, all marsh, all woodland, yields its particular set of birds,
perhaps in profusion \ but the kinds will be limited in number. It
is of first importance to remember this, when you are so fortunate
as to have choice of a collecting-ground ; and it will guide your
steps aright in a day’s walk anywhere, for it will make you leave
covert for open, wet for dry, high for low, and back again. Well-
watered country is more fruitful of bird-life than desert or prairie ;
warm regions are more productive than cold ones. As a rule,
variety and abundance of birds are in direct ratio to diversity and
luxuriance of vegetation. Your most valuable as well as largest
bags may be made in the regions most favoured botanically, up to
the point where exuberance of plant -growth mechanically opposes
your operations.

Search for particular Birds can only be well directed by a
knowledge of their special haunts and habits, and is one of the
mysteries of wood-craft to be solved by long experience and close
observation. Here is where the true naturalist bears himself with
conscious pride and strength, winning laurels that become him, and
do honour to his calling. Where to find game (“game ” is anything
that vulgar people do not ridicule you for shooting) of all the kinds
we have in this country has been so often and so minutely detailed
in sporting-works that it need not be here enlarged upon, especially
since, being the best known, game-birds are the least valuable of
ornithological material. Most large or otherwise conspicuous birds
have very special haunts that may be soon learned ; and as a rule
such rank next after game in ornithological disesteem. Birds of
prey are an exception to these statements ; they range everywhere,
and most of them are worth securing. Hawks will unwittingly fly
in your way oftener than they will allow you to approach them
when perched : be ready for them. Owls will be startled out of
their retreats in thick bushes, dense foliage, and hollow trees, in the
daytime ; if hunting them at night, good aim in the dark may be
taken by rubbing a wet lucifer match on the sight of the gun,
causing a momentary glimmer. Large and small waders are to be
found by any water’s edge, in open marshes, and often on dry

SEC. Ill SUGGESTIONS AND DIRECTIONS FOR FIELD-WORK 17

plains; the herons more particularly in heavy bogs and dense
swamps. Under cover, waders are oftenest approached by stealth ;
in the open, by strategy ; but most of the smaller kinds require the
exercise of no special precautions. Swimming - birds, aside from
water-fowl (as the “game” kinds are called), are generally shot
from a boat, as they fly past ; but at their breeding-places many
kinds that congregate in vast numbers are readily reached. There
is a knack of shooting loons and grebes on the water ; if they are
to be reached at all by the shot it will be by aiming not directly at
them but at the water just in front of them. They do not go under
just where they float, but kick up behind like a jumping-jack and
plunge forward. Rails and several kinds of sparrows are confined
to reedy marshes. But why prolong such desultory remarks ?
Little can be said to the point without at least a miniature treatise
on ornithology ; and I have not yet even alluded to the diversified
host of small insectivorous and granivorous birds that fill our woods
and fields. The very existence of most of these is unknown to all
but the initiated ; yet they include the treasures of the orni-
thologist. Some are plain and humble, others are among the most
beautiful objects in nature ; but most agree in being small, and
therefore liable to be overlooked. The sum of my advice about
them must be brief. Get over as much ground, both wooded and
open, as you can thoroughly examine in a day’s tramp, and go out
as many days as you can. It is not always necessary, however, to
keep on the tramp, especially during the migration of the restless
insectivorous species. One may often shoot for hours without
moving more than a few yards, by selecting a favourable locality
and allowing the birds to come to him as they pass in varied troops
through the low woodlands or swampy thickets. Keep your eyes
and ears wide open. Look out for every rustling leaf and swaying
twig and bending blade of grass. Hearken to every note, however
faint ; when there is no sound, listen for a chirp. Habitually move
as noiselessly as possible. Keep your gun always ready. Improve
every opportunity of studying a bird you do not wish to destroy ;
you may often make observations more valuable than the specimen.
Let this be the rule with all birds you recognise. But I fear I
must tell you to shoot an unknown bird on sight ; it may give you
the slip in a moment and a prize may be lost. One of the most
fascinating things about field-work is its uncertainty ; you never
know what’s in store for you as you start out ; you never can tell
what will happen next ; surprises are always in order, and excite-
ment is continually whetted on the chances of the varied chase.

For myself, the time is past, happily or not, Avhen every bird
was an agreeable surprise, for dewdrops do not last all day ; but I
have never yet walked in the woods without learning something
C

i8

FIELD ORNITHOLOGY

PART I

pleasant that I did not know before. I should consider a bird new
to science ample reward for a month’s steady work ; one bird new
to a locality would repay a week’s search ; a day is happily spent that
shows me any bird that I never saw alive before. How then can you,
with so much before you, keep out of the woods another minute %
All Times are good times to go a -shooting ; but some
are better than others. {a) Time of year. In all temperate
latitudes, spring and fall — periods of migration with most birds —
are the most profitable seasons for collecting. Not only are birds
then most numerous, both as species and as individuals, and most
active, so as to be the more readily found, but they include a far
larger proportion of rare and valuable kinds. In every locality in
this country the periodical visitants outnumber the permanent
residents ; in most regions the number of regular migrants, that
simply pass through in the spring and fall, equals or exceeds that
of either of the sets of species that come from the south in spring
to breed during the summer, or from the north to spend the winter.
Far north, of course, on or near the limit of the vernal migration,
where there are few if any migrants, and where the winter birds
are extremely few, nearly all the bird-fauna is composed of “ summer
visitants ” ; far south the reverse is somewhat the case, though with
many qualifications. Between these extremes, what is conventionally
known as “ a season ” means the period of the vernal or autumnal
migration. Look out, then, for “ the season,” and work all through
it at a rate you could not possibly sustain the year around. (f)
Time of day. Early in the morning and late in the afternoon are
the best times for birds. There is a mysterious something in these
diurnal crises that sets bird-life astir, over and above what is explain-
able by the simple fact that they are the transition periods from
repose to activity, or the reverse. Subtle meteorological changes
occur ; various delicate instruments used in physicists’ researches
are sometimes inexplicably disturbed ; diseases have often their
turning point for better or worse ; people are apt to be born or
die ; and the susceptible organisms of birds manifest various excite-
ments. Whatever the operative influence, the fact is, birds are
particularly lively at such hours. In the dark they rest — most of
them do ; at noonday, again, they are comparatively still ; between
these times they are passing to or from their feeding grounds or
roosting places ; they are foraging for food, they are singing ; at
any rate, they are in motion. Many migratory birds (among them
warblers, etc.) perform their journeys by night; just at daybreak
they may be seen to descend from the upper regions, rest a while,
and then move about briskly, singing and searching for food. Their
meal taken, they recuperate by resting till towards evening ; feed
again and are off for the night. If you have had some experience,

SEC. Ill SUGGESTIONS AND DIRECTIONS FOR FIELD' WORN 19

don’t you remember what a fine spurt you made early that morning ?
— how many unexpected shots offered as you trudged home belated
that evening ? Now I am no fowl, and have no desire to adopt
the habits of the hen-yard ; I have my opinion of those who like
the world before it is aired ; I think it served the worm right for
getting up, when caught by the early bird ; nevertheless I go shoot-
ing betimes in the morning, and would walk all night to find a rare
bird at daylight, (c) Weather. It rarely occurs in this country
that either heat or cold is unendurably severe ; but extremes of
temperature are unfavourable, for two reasons : they both occasion
great personal discomfort ; and in one extreme only a few hardy
birds will be found, while in the other most birds are languid, dis-
posed to seek shelter, and therefore less likely to be found. A
still, cloudy day of moderate temperature offers as a rule the best
chance ; among other reasons, there is no sun to blind the eyes, as
always occurs on a bright day in one direction, particularly when
the sun is low. While a bright day has its good influence in set-
ting many birds astir, some others are most easily approached in
heavy or falling weather. Some kinds are more likely to be secured
during a light snowfall, or after a storm. Singular as it may seem,
a thoroughly wet day offers some peculiar inducements to the col-
lector. I cannot well specify them, but I heartily indorse a remark
John Cassin once made to me: “I like,” said he, “to go shooting
in the rain sometimes ; there are some curious things to be learned
about birds when the trees are dripping ; things, too, that have not
yet found their way into the books.”

How many Birds of the Same Kind do you want ? —
All you can get — with some reasonable limitations ; say fifty or a
hundred of any but the most abundant and widely diffused species.
You may often be provoked with your friend for speaking of some
bird he shot, but did not bring you, because, he says, “ Why, you’ve
got one like that ! ” Birdskins are capital ; capital unemployed
may be useless, but can never be worthless. Birdskins are a medium
of exchange among ornithologists the world over ; they represent
value — money value and scientific value. If you have more of one
kind than you can use, exchange with some one for species you
lack ; both parties to the transaction are equally benefited. Let
me bring this matter under several heads, {a) Your own series
of skins of any species is incomplete until it contains at least one
example of each sex, of every normal state of plumage, and every
normal transition stage of plumage, and further illustrates the
principal abnormal variations in size, form, and colour to which
the species may be subject j I will even add that every different
faunal area the bird is known to inhabit should be represented by
a specimen, particularly if there be anything exceptional in the

20

FIELD ORNITHOLOGY

PART I

geographical distribution of the species. Any additional specimens
to all such are your only “ duplicates, ” properly speaking. (6)
Birds vary so much in their size, form, and colouring, that a “ spe-
cific character ” can only be precisely determined from examination
of a large number of specimens, shot at different times, in different
places ; still less can the “ limits of variation ” in these respects be
settled without ample materials, (c) The rarity of any bird is an
arbitrary and fluctuating consideration, because in the nature of the
case there can be no natural unit of comparison, nor standard of
appreciation. It may be said, in general terms, no bird is actually
‘‘ rare.” With a few possible exceptions, as in the cases of birds
occupying extraordinarily limited areas, like some of the birds-of-
paradise, or about to become extinct, like the pied duck,^ enough
birds of all kinds exist to overstock every public and private collec-
tion in the world, without sensible diminution of their numbers.
“ Karity ” or the reverse is only predicable upon the accidental (so
to speak) circumstances that throw, or tend to throw, specimens
into naturalists’ hands. Accessibility the variable element in every
case. The fulmar petrel ^ is said (on what authority I know not) to
exceed any other bird in its aggregate of individuals ; how do the
skins of that bird you have handled compare in number with speci-
mens you have seen of the “rare” warbler of your own vicinity?
All birds are common somewhere at some season : the point is, have
collectors been there at the time ? Moreover, even the arbitrary
appreciation of “rarity” is fluctuating, and may change at any
time; long- sought and highly -prized birds are liable to appear
suddenly in great numbers in places that knew them not before ;
a single heavy invoice of a bird from some distant or little-
explored region may at once stock the market, and depreciate the
current value of the species to almost nothing. For example,
Baird’s bunting^ and Sprague’s lark‘d remained for thirty years
among special desiderata, only one specimen of the former and two
or three of the latter being known. Yet they are two of the most
abundant birds of Dakota, where in 1873 I took as many of both
as I desired ; and specimens enough have lately been secured to
stock all the leading museums of both Europe and America, {d)
Some practical deductions are to be made from these premisses.
Your object is to make yourself acquainted with all the birds of
your vicinity, and to preserve a complete suite of specimens of
every species. Begin by shooting every bird you can, coupling
this sad destruction, however, with the closest observations upon
habits. You will very soon fill your series of a few kinds, that
you find almost everywhere, almost daily. Then if you are in a

^ Camptolcemus Idbradorius. ^ Fulmarus glacialis.

^ Passer cuius {Centronyx) hairdi. Anthus {Neocorys) spraguei.

SEC. Ill SUGGESTIONS AND DIRECTIONS EOR EIELD-WORK 2:

region the ornithology of which is well known, at once stop killing
these common birds — they are in every collection. Keep an eye
on them, studying them always, but turn your actual pursuit into
other channels, until in this way, gradually eliminating the un-
desirables, you exhaust the bird-fauna as far as possible (you will not
quite exhaust it — at least for many years). But if you are in a new
or little-known locality, I had almost said the very reverse course
is the best. The chances are that the most abundant and character-
istic birds there are “ rare ” in collections. Many a bird’s range
is quite restricted : you may happen to be just at its metropolis ;
seize the opportunity, and get good store, — yes, up to fifty or a
hundred ; all you can spare will be thankfully received by those who
have none. Quite as likely, birds that are scarce just where you
happen to be, are so only because you are on the edge of their
habitat, and are plentiful in more accessible regions. But, rare or
not, it is always a point to determine the exact geographical dis-
tribution of a species ; and this is fixed best by having specimens
to tell each its own tale, from as many different and widely-separated
localities as possible. This alone warrants procuring one or more
specimens in every locality ; the commonest bird acquires a certain
value if it be captured away from its ordinary range. But let all
your justifiable destruction of birds be tempered with mercy j your
humanity will be continually shocked with the havoc you work,
and should never permit you to take life wantonly. Never shoot
a bird you do not fully intend to preserve, or to utilise in some
proper way. Bird -life is too beautiful a thing to destroy to no
purpose; too sacred a thing, like all life, to be sacrificed, unless
the tribute is hallowed by worthiness of motive. “Not a sparrow
falleth to the ground without His notice.”

I should not neglect to speak particularly of the care to be taken
to secure full suites of females. Most miscellaneous collections con-
tain four or more males to every female — a disproportion that should
be as far reduced as possible. The reason for this disparity is obvious :
females are usually more shy and retiring in disposition, and less
frequently noticed ; while their smaller size and plainer plumage, as
a rule, further favour their concealment. The difference in colour-
ing is greatest among those groups where the males are most richly
clad, and the shyness of the mother birds is most marked during
the breeding season, just when the males, full of song, and in their
nuptial attire, become most conspicuous. It is often worth while
to neglect the gay Benedicts, to trace out and secure the plainer but
not less interesting females. This pursuit, moreover, often leads to
discovery of the nests and eggs, — an important consideration.
Although both sexes are generally found together when breeding,
and mixing indiscriminately at other seasons, they often go in sepa-

22

FIELD ORNITHOLOGY

PART I

rate flocks, and often migrate independently of each other ; in this
case the males usually in advance. Towards the end of the passage
of some warblers, for instance, we may get almost nothing hut
females, all our specimens of a few days before having been males.
The notable exceptions to the rule of smaller size of the female are
among rapacious birds and many waders, though in these last the
disparity is not so marked. I only recall one instance, among
English birds, of the female being more richly coloured than the
male — the phalaropes. When the sexes are notably different in
adult life, the young of both sexes usually resemble the adult female,
the young males gradually assuming their distinctive characters.
When the adults of both sexes are alike, the young commonly differ
from them.

In the same connection I wish to urge a point, the importance
of which is often overlooked ; it is our practical interpretation of
the adage, “ a bird in the hand is worth two in the bush.” Always
keep the first specimen you secure of a species till you get another,
no matter how common the species, how poor the specimen, or
how certain you may feel of getting others. Your most reason-
able calculations may come to naught, from a variety of circum-
stances, and any specimen is better than no specimen, on general
principles. And in general, do not, if you can help it, discard any
specimen in the field. No tyro can tell what will prove valuable
and what not ; while even the expert may regret to find that a
point comes up which a specimen he injudiciously discarded might
have determined. Let a collection be “ weeded out,” if at all, only
after deliberate and mature examination, when the scientific results
it affords have been elaborated by a competent ornithologist ; and
even then, the refuse (with certain limitations) had better be put
where it will do some good, than be destroyed utterly. If forced to
reduce bulk, owing to limited facilities for transportation in the field
(as too often happens), throw away according to size, other things
being equal. Given only so many cubic inches or feet, eliminate
the few large birds which take up the space that would contain
fifty or a hundred different little ones. If you have a fine large
eagle or pelican, for instance, throw it away first, and follow it with
your ducks, geese, etc. In this way, the bulk of a large miscel-
laneous collection may be reduced one-half, perhaps, with very little
depreciation of its actual value. The same principle may be
extended to other collections in natural history (excepting fossils,
which are always weighty, if not also bulky) ; very few birdskins,
indeed, being as valuable contributions to science as, for example, a
vial of insects that occupies no more room may prove to be.

What is “ A Good Day’s Work ” ? — Fifty birds shot, their skins
preserved, and observations recorded, is a very good day’s work ; it

SEC. Ill SUGGESTIONS AND DIRECTIONS FOR FIELD- WORK 23

is sharp practice, even when birds are plentiful. I never knew a
person to average anywhere near it ; even during the “ season ”
such work cannot possibly be sustained. You may, of course, by a
murderous discharge into a flock, get a hundred or more in a
moment ; but I refer to collecting a fair variety of birds. You will
do very well if you average a dozen a day during the seasons. I
doubt whether any collector ever averaged as many the year around ;
it would be over four thousand specimens annually. The greatest
number I ever procured and prepared in one day was forty, and I
have not often gone over twenty. Even when collecting regularly
and assiduously, I am satisfied to average a dozen a day during the
migrations, and one-third or one-fourth as many the rest of the
year. Probably this implies the shooting of about one in five not
skinned for various reasons, as mutilation, decay, or want of time.

Approaching Birds. — There is little if any trouble in getting near
enough to shoot most birds. With notable exceptions, they are
harder to see when near enough, or to hit when seen j particularly
small birds that are almost incessantly in motion. As a rule — and
a curious one it is — difficulty of approach is in direct ratio to the
8%ze of the bird ; it is perhaps because large conspicuous birds are
objects of more general pursuit than the little ones you ordinarily
search for. The qualities that birds possess for self-preservation
may be called wanness in large birds, shyness in small ones. The
former make off knowingly from a suspicious object j the latter fly
from anything that is strange to them, be it dangerous or not. This
is strikingly illustrated in the behaviour of small birds in the
wilderness, as contrasted with their actions about towns ; they are
more timid under the former circumstances than when grown
accustomed to the presence of man. It is just the reverse with a
hawk or raven, for instance in populous districts they spend much
of their time in trying to save their skins, while in a new country
they have not learned, like Indians, that a white man is “mighty
uncertain.” In stealing on a shy bird, you will of course take ad-
vantage of any cover that may offer, as inequalities of the ground,
thick bushes, the trunks of trees ; and it is often worth while to
make a considerable d4tour to secure unobserved approach. I think
that birds are more likely, as a rule, to be frightened away by the
movements of the collector, than by his simple presence, however
near, and that they are more afraid of noise than of mere motion.
Crackling of twigs and rustling of leaves are sharp sounds, though
not loud ones ; you may have sometimes been surprised to find how
distinctly you could hear the movements of a horse or cow in
underbrush at some distance. Birds have sharp ears for such
sounds. Form a habit of stealthy movement ; it tells, in the long
run, in comparison with lumbering tread. There are no special

24

FIELD ORNITHOLOGY

PART I

precautions to be taken in shooting through high open forest ; you
have only to saunter along with your eyes in the tree-tops. It is
ordinarily the easiest and on the whole the most remunerative path
of the collector. In traversing fields and meadows move briskly,
your principal object being to flush birds out of the grass j and as
most of your shots will be snap ones, keep in readiness for instant
action. Excellent and varied shooting is to be had along the
hedgerows, and in the rank herbage that fringes fences. It is best
to keep at a little distance, yet near enough to arouse all the
birds as you pass ; you may catch them on wing, or pick them off
just as they settle after a short flight. In this shooting, two
persons, one on each side, can together do more than twice as
much work as one. Thickets and tangled undergrowth are
favourite resorts of many birds ; but when very close, or, as often
happens, over miry ground, they are hard places to shoot in. As
you come thrashing through the brush, the little inhabitants are
scared into deeper recesses ; but if you keep still a few minutes in
some favourable spot, they are reassured, and will often come back
to take a peep at you. A good deal of standing still will repay you
at such times ; needless to add, you cannot be too lightly loaded for
such shooting, when birds are mostly out of sight if a dozen yards
off. When yourself concealed in a thicket, and no birds appear,
you can often call numbers about you by a simple artifice. Apply
the back of your hand to your slightly parted lips, and suck in air;
it makes a nondescript screeping noise, variable in intonation at
your whim, and some of the sounds resemble the cries of a wounded
bird, or a young one in distress. It wakes up the whole neigh-
bourhood, and sometimes puts certain birds almost beside them-
selves, particularly in the breeding season. Torturing a wounded
bird to make it scream in agony accomplishes the same result, but
of course is only permissible under great exigency. In penetrating
swamps and marshes, the best advice I can give you is to tell you
to get along the best way you can. Shooting on perfectly open
ground offers much the same case ; you must be left to your, own
devices. I will say, however, you can ride on horseback, or even
in a buggy, nearer birds than they will allow you to walk up to
them. Sportsmen take advantage of this to get within a shot of
the upland plover, usually a very wary bird in populous districts ;
I have driven right into a flock of wild geese ; in California they
often train a bullock to graze gradually up to geese, the gunner
being hidden by its body. There is one trick worth knowing ; it
is not to let a bird that has seen you know by your action that
you have seen it, but to keep on unconcernedly, gradually sidling
nearer. I have secured many hawks in this way, when the bird
would have flown off at the first step of direct approach. Number-

SEC. Ill SUGGESTIONS AND DIRECTIONS FOR FIELD-WORK 25

less other little arts will come to you as your wood - craft
matures.

Recovering Birds. — It is not always that you secure the birds
you kill ; you may not he able to find them, or you may see them
lying, perhaps but a few feet off, in a spot practically inaccessible.
Under such circumstances a retriever does excellent service, as
already hinted; he is equally useful when a bird properly ‘‘marked
down is not found there, having fluttered or run away and hidden
elsewhere. The most difficult of all places to find birds is among
reeds, the sameness of which makes it almost impossible to redis-
cover a spot whence the eye has once wandered, while the peculiar
growth allows birds to slip far down out of sight. In rank grass or
weeds, when you have walked up with your eye fixed on the spot
where the bird seemed to fall, yet failed to discover it, drop your
cap or handkerchief for a mark, and hunt around it as a centre, in
enlarging circles. In thickets, make a bee-line for the spot, if
possible keeping your eye on the spray from which the bird fell, and
not forgetting where you stood on firing ; you may require to come
back to the spot and take a new departure. You will not seldom
see a bird just shot at fly off as if unharmed, when really it will
drop dead in a few moments. In all cases, therefore, when the
bird does not drop at the shot, follow it with your eyes as far as
you can ; if you see it finally drop, or even flutter languidly down-
ward, mark it on the principles just mentioned, and go in search.
Make every endeavour to secure wounded birds, on the score of
humanity ; they should not be left to pine away and die in linger-
ing misery if it can possibly be avoided.

Killing Wounded Birds. — You will often recover winged birds,
as full of life as before the bone was broken ; and others too griev-
ously hurt to fly, j^et far from death. Your object is to kill them
as quickly and painlessly as possible, without injuring the plumage.
This is to be accomplished, with all small birds, by suffocation.
The respiration and circulation of birds is very active, and most of
them die in a few moments if the lungs are so compressed that
they cannot breathe. Squeeze the bird tightly across the chest,
under the wings, thumb on one side, middle finger on the other,
forefinger pressed in the hollow at the root of the neck, between
the forks of the merrythought. Press firmly, hard enough to fix
the chest immovably and compress the lungs, but not to break in
the ribs. The bird will make vigorous but ineffectual efforts to
breathe, when the muscles will contract spasmodically ; but in a
moment more, the system relaxes with a painful shiver, light fades
from the eyes, and the lids close. I assure you, it will make you
wince the first few times ; you had better hold the poor creature
behind you. You can tell by its limp feel and motionlessness when

26

FIELD ORNITHOLOGY

PART I

it is dead, without watching the sad struggle. Large birds cannot
be dealt with in this way ; I would as soon attempt to throttle a
dog as a loon, for instance, upon which all the pressure you can
give makes no sensible impression. A winged hawk, again, will
throw itself on its back as you come up, and show such good fight
with beak and talons, that you may be quite severely scratched in
the encounter : meanwhile the struggling bird may be bespattering
its plumage with blood. In such a case — in any case of a large
bird making decided resistance — I think it best to step back a few
paces and settle the matter with a light charge of mustard-seed.
Any large bird once secured may be speedily despatched by stabbing
to the heart with some slender instrument thrust in under the
wing — care must be taken too about the bleeding ; or, it may be
instantly killed by piercing the brain with a knife introduced into
the mouth and driven upward and obliquely backward from the
palate. The latter method is preferable, as it leaves no outward
sign and causes no bleeding to speak of. With your thumb, you
may indent the back part of a small bird’s skull so as to compress
the cerebellum, which causes instant death. It is useless to compress
the windpipe of a bird whose wing is broken near the shoulder, for
the bone is hollow, and the bird can breathe through it.

Handling* Bleeding Birds. — Bleeding depends altogether upon
what part or organ is wounded; but, other things being equal,
violence of the haemorrhage is usually in direct proportion to the
size of the shot -hole; when mustard -seed is used it is ordinarily
very trifling, if it occur at all. Blood flows oftener from the orifice
of exit of a shot, than from the wound of entrance, for the latter is
usually plugged with a little wad of feathers. Bleeding from the
mouth or nostrils is the rule when the lungs are wounded. When
it occurs, hold up the bird by the feet, and let it drip ; a general
squeeze of the body in that position will facilitate the drainage.
In general, hold a bird so that a bleeding place is most dependent ;
then, pressure about the part will help the flow. A “ gob ” of
blood, which is simply a forming clot, on the plumage may often
be dexterously flipped almost clean away with a snap of the finger.
It is first-rate practice to take cotton and forceps into the field to
plug up shot-holes, and stop the mouth and nostrils and vent on
the spot. I follow the custom of the books in recommending this,
but I suspect that only a few of the most leisurely and elegant
collectors do so habitually. Shot-holes may be found by gently
raising the feathers, or blowing them aside ; you can of course get
only a tiny plug into the wound itself, but it should be one end of
a sizable pledget, the rest lying fluffy among the feathers. In
stopping the mouth or vent, ram the fluff of cotton entirely inside.
You cannot conveniently stop up the nostrils of small birds sepa-

SEC. Ill SUGGESTIONS AND DIRECTIONS FOR FIEID-WORK 27

rately ; but take a light cylinder of cotton, lay it transversely across
the base of the upper mandible, closely covering the nostrils, and
confine it there by tucking each end tightly into the corner of the
mouth. In default of such nice fixing as this, a pinch of dry loam
pressed on a bleeding spot will plaster itself there and stop further
mischief. Never try to wi]^e q/f fresh blood that has already wetted
the plumage ; you will only make matters worse. Let it dry on,
and then — but the treatment of blood-stains, and other soilings of
plumage, is given beyond.

Carrying Birds Home Safe. — Suppose you have secured a fine
specimen, very likely without a soiled or ruffled feather ; your next
care will be to keep it so till you are ready to skin it. But if you
pocket or bag it directly, it will be a sorry-looking object before
you get home. Each specimen must be separately cared for, by
wrapping in stout paper ; writing paper is as good as any, if not
the best. It will repay you to prepare a stock of paper before
starting out ; your most convenient sizes are those of a half-sheet
of note, of letter, and of cap respectively. Either take these, or
fold and cut newspaper to correspond. Plenty of paper will go in
the breast pockets of the shooting coat. Make a “cornucopia,” —
the simplest thing in the world, but, like tying a particular knot,
hard to explain. Setting the wings closely, adjusting disturbed
feathers, and seeing that the bill points straight forward, thrust the
bird head-ffrst into one of these paper cones, till it will go no
farther, being bound by the bulge of the breast. Let the cone be
large enough for the open end to fold over or pinch together entirely
beyond the tail. Be particular not to crumple or bend the tail-
feathers. Lay the paper cases in the game bag or great pocket so
that they very nearly run parallel and lie horizontal j they will
carry better than if thrown in at random. Avoid overcrowding the
packages, as far as is reasonably practicable ; moderate pressure
will do no harm, but if great it may make birds bleed afresh, or
cause the fluids of a wounded intestine to ooze out and soak the
plumage of the belly, — a very bad accident indeed. For similar
obvious reasons, do not put a large heavy bird on top of a lot of
little ones ; I would sooner sling a hawk or heron over my shoulder,
or carry it by hand. If it goes in the bag, see that it gets to the
bottom. Avoid putting birds in pockets that are close about your
person \ they are almost always unduly pressed, and may gain
enough additional warmth from your body to make them begin to
decompose before you are ready to skin them. Handle birds no
more than is necessary, especially white-plumaged ones j ten to one
your hands are powder-begrimed : and besides, even the warmth
and moisture of your palms may tend to injure a delicate feather-
ing. Ordinarily pick up a bird by the feet or bill ; as you need

28

FIELD ORNITHOLOGY

PART I

both hands to make the cornucopia, let the specimen dangle by the
toes from your teeth while you are so employed. In catching at a
wounded bird, aim to cover it entirely with your hand ; but what-
ever you do, never seize it by the tail, which then will often be
left in your hands for your pains. Never grasp wing-tips or tail-
feathers ; these large flat quills would get a peculiar crimping all
along the webs, very difficult to efface. Finally, I would add, there
is a certain knack or art in manipulating, either of a dead bird or
a birdskin, by which you may handle it with seeming carelessness
and perfect impunity ; whilst the most gingerly fingering of an
inexperienced person will leave its rude trace. You will naturally
acquire the correct touch ; but it can be neither taught nor described.
While the ordinary run of land-birds will be brought home in good
order by the foregoing method, some require special precautions.
I refer to sea-birds, such as gulls, terns, petrels, etc., shot from a
boat. In the first place, the plumage of most of them is, in part
at least, white and of exquisite purity. Then, fish-eating birds
usually vomit and purge when shot. They are necessarily fished
all dripping from the water. They are too large for pocketing.
If you put them on the thwarts or elsewhere about the boat, they
usually fall off, or are knocked off, into the bilge water ; if you stow
them in the cubby-hole, they will assuredly soil by mutual pressure,
or by rolling about. It will repay you to pick them from the
water by the bill, and shake off all the water you can ; hold them
up, or let some one do it, till they are tolerably dry ; plug the
mouth, nostrils, and vent, if not also shot -holes; wrap each one
separately in a cloth (not paper) or a mass of tow, and pack steadily
in a covered box or basket taken on board for this purpose. With
such precautions as these, birds most liable to be soiled reach the
skinning-table in perfect order ; and your care will afterward trans-
form them into specimens without spot or blemish.

§ 4.— HYGIENE OF COLLECTOESHIP

It is unnecessary to speak of the Healthfulness of a pursuit
that, like the collector’s occupation, demands regular bodily exercise,
and at the same time stimulates the mind by supplying an object,
thus calling the whole system into exhilarating action. Yet collect-
ing has its perils, not to be overlooked if we would adequately
guard against them, as fortunately we may, in most cases, by simple
precautions. The dangers of taxidermy itself are elsewhere noticed ;
but, besides these, the collector is exposed to vicissitudes of the
weather, may endure great fatigue, may breathe miasm, and may
be mechanically injured.

SEC. IV

HYGIENE OF COLLECTORSHIP

29

Accidents from the Gun have been already noticed ; a few
special rules will render others little liable to occur. The secret of
safe climling is never to relax one hold until another is secured ; it
is equally applicable to scrambling over rocks, a particularly difficult
thing to do safely with a loaded gun. Test rotten, slippery, or
otherwise suspicious holds before trusting them. In lifting the
body up anywhere, keep the mouth shut, breathe through the
nostrils, and go slowly. In swimming, waste no strength unneces-
sarily in trying to stem a current ; yield partly, and land obliquely
lower down ; if exhausted, float ; the slightest motion of the hands
will ordinarily keep the face above water ; and in any event keep
your wits collected. In fording deeply, a heavy stone will strengthen
your position. Never sail a boat experimentally; if you are no
sailor, take one with you or stay on land. In crossing a high,
narrow footpath, never look lower than your feet ; the muscles will
work true if not confused with faltering instructions from a giddy
brain. On soft ground, see what, if anything, has preceded you ;
large hoof-marks generally mean that the way is safe ; if none are
found, inquire for yourself before going on. Quicksand is the most
treacherous, because far more dangerous than it looks. Cattle-
paths, however erratic, commonly prove the surest way out of a
difficult place, whether of uncertain footing or dense undergrowth.

Miasm. — Unguarded exposure in malarious regions usually
entails sickness, often preventable, however, by due precautions. It
is worth knowing, in the first place, that miasmatic poison is most
powerful between sunset and sunrise ; more exactly, from the damp
of the evening until night-vapours are dissipated ; we may be out
in the daytime with comparative impunity, where to pass a night
would be almost certain disease. If forced to camp out, seek the
highest and driest spot, put a good fire on the swamp side, and
also, if possible, let trees intervene. Never go out on an empty
stomach; just a cup of coffee and a crust may make a decided
difference. Meet the earliest unfavourable symptoms with quinine;
I should rather say, if unacclimated, anticipate them with this
invaluable agent. Endeavour to maintain high health of all
functions by the natural means of regularity and temperance in diet,
exercise, and repose.

“Taking Cold.” — This vague “ household word ” indicates one
or more of a long varied train of unpleasant affections, nearly
always traceable to one or the other of only two causes ; sudden
change of temperature, and unequal distribution of temperature on
the surface of the person. No extremes of heat or cold can alone
effect this result ; persons frozen to death do not “ take cold ”
during the process. But if a part of the body be rapidly cooled, as
by evaporation from a wet article of clothing, or by sitting in a

30

FIELD ORNITHOLOGY

PART I

draught of air, the rest of the surface remaining at an ordinary
temperature ; or if the temperature of the whole he suddenly
changed by going out into the cold, or by coming into a warm room,
there is much liability of trouble. There is an old saying —

When the air comes through a hole

Say your prayers to save your soul ;

and I should think almost any one could get “a cold” with a
spoonful of water on the wrist held to a key-hole. Singular as it
may seem, sudden warming when cold is more dangerous than the
reverse; every one has noticed how soon the handkerchief is
required on entering a heated room on a cold day. Frost-bite offers
an extreme illustration of this. As the Irishman said on picking
himself up, it was not the fall, but stopping so quickly, that hurt
him ; it is not the gradual lowering of the temperature to the
freezing-point, but its subsequent sudden elevation, that devitalises
the tissue. This is why rubbing with snow, or bathing in cold
water, is required to restore safely a frozen part ; the arrested
.circulation must be very gradually re-established, or inflammation,
perhaps mortification, ensues. General precautions against taking
cold are almost self-evident in this light. There is ordinarily little
if any danger to be apprehended from wet clothes, so long as
exercise is kept up ; for the glow compensates for the extra cooling
by evaporation. Nor is a complete drenching more likely to be
injurious than wetting of one part. But never sit still wet ; and
in changing rub the body dry. There is a general tendency,
springing from fatigue, indolence, or indifference, to neglect damp
feet ; that is to say, to dry them by the fire ; but this process is
tedious and uncertain. I would say especially, off with the muddy
boots and sodden socks at once ; dry stockings and slippers, after a
hunt, may make just the difference of your being able to go out
again or never. Take care never to check perspiration; during
this process, the body is in a somewhat critical condition, and
sudden arrest of the function may result disastrously, even fatally.
One part of the business of perspiration is to equalise bodily
temperature, and it must not be interfered with. The secret of
much that might be said about bathing when heated lies here. A
person overheated, panting it may be, with throbbing temples, and
a dry skin, is in danger partly because the natural cooling by
evaporation from the skin is denied, and this condition is sometimes
not far from a sunstroke. Under these circumstances, a person of
fairly good constitution may plunge into the water with impunity,
even with benefit. But if the body be already cooling by sweating,
rapid abstraction of heat from the surface may cause internal
congestion, never unattended with danger. Drinking ice -water

SEC. IV

HYGIENE OF COLLECTORSHIP

31

offers a somewhat parallel case ; even on stooping to drink at the
brook, when flushed with heat, it is well to bathe the face and
hands first, and to taste the water before a full draught. It is a
well-known excellent rule, not to bathe immediately after a full
meal ; because during digestion the organs concerned are compara-
tively engorged, and any sudden disturbance of the circulation may
be disastrous. The imperative necessity of resisting drowsiness
under extreme cold requires no comment. In walking under a hot
sun, the head may be sensibly protected by green leaves or grass in
the hat; they may be advantageously moistened, but not enough
to drip about the ears. Under such circumstances the slightest
giddiness, dimness of sight, or confusion of ideas, should be taken
as a warning of possible sunstroke, instantly demanding rest and
shelter.

Hunger and Fatigue are more closely related than they
might seem to be ; one is a sign that the fuel is out, and the other
asks for it. Extreme fatigue, indeed, destroys appetite ; this simply
means, temporary incapacity for digestion. But even far short of
this, food is more easily digested and better relished after a little
preparation of the furnace. On coming home tired, it is much
better to make a leisurely and reasonably nice toilet than to eat at
once, or to sit still thinking how tired you are ; after a change and
a wash you will feel like a “ new man,” and go to table in capital
state. Whatever dietetic irregularities a high state of civilisation
may demand or render practicable, a normally healthy person is
inconvenienced almost as soon as his regular meal-time passes with-
out food ; a few can work comfortably or profitably fasting over six
or eight hours. Eat before starting ; if for a day’s tramp, take a
lunch ; the most frugal meal will appease if it do not satisfy hunger,
and so postpone its urgency. As a small scrap of practical wisdom,
I would add, keep the remnants of the lunch, if there are any ; for
you cannot always be sure of getting in to supper.

Stimulation. — When cold, fatigued, depressed in mind, and on
other occasions, you may feel inclined to resort to artificial stimulus.
Eespecting this many-sided theme I have a few words to offer of
direct bearing on the collector’s case. It should be clearly under-
stood in the first place that a stimulant confers no strength what-
ever ; it simply calls the powers that be into increased action at
their own expense. Seeking real strength in stimulus is as wise as
an attempt to lift yourself up by the boot-straps. You may gather
yourself to leap the ditch and you clear it ; but no such muscular
energy can be sustained ; exhaustion speedily renders further ex-
penditure impossible. But now suppose a very powerful mental
impression be made, say the circumstance of a succession of ditches
in front, and a mad dog behind ; if the stimulus of terror be suffi-

32

FIELD ORNITHOLOGY

PART I

ciently strong, you may leap on till you drop senseless. Alcoholic
stimulus is a parallel case, and is not seldom pushed to the same
extreme. Under its influence you never can tell when you are
tired j the expenditure goes on, indeed, with unnatural rapidity,
only it is not felt at the time ; but the upshot is you have all the
original fatigue to endure and to recover from, plus the fatigue
resulting from over-excitation of the system. Taken as a fortification
against cold, alcohol is as unsatisfactory as a remedy for fatigue.
Insensibility to cold does not imply protection. The fact is the
exposure is greater than before ; the circulation and respiration
being hurried, the waste is greater, and as sound fuel cannot be
immediately supplied, the temperature of the body is soon lowered.
The transient warmth and glow over, the system has both cold and
depression to endure ; there is no use in borrowing from yourself
and fancying you are richer. Secondly, the value of any stimulus
(except in a few exigencies of disease or injury) is in proportion,
not to the intensity, but to the equableness and durability of its
effect. This is one reason why tea, coffee, and articles of corre-
sponding qualities are preferable to alcoholic drinks ; they work so
smoothly that their effect is often unnoticed, and they “ stay by ”
well ; the friction of alcohol is tremendous in comparison. A glass
of grog may help a veteran over the fence, but no one, young or
old, can shoot all day on liquor. I have had so much experience in
the use of tobacco as a mild stimulant that I am probably no impartial
judge of its merits : I will simply say I do not use it in the field,
because it indisposes to muscular activity, and favours reflection
when observation is required ; and because temporary abstinence
provokes the morbid appetite and renders the weed more grateful
afterwards. Thirdly, undue excitation of any physical function is
followed by corresponding depression, on the simple principle that
action and reaction are equal ; and the balance of health turns too
easily to be wilfully disturbed. Stimulation is a draft upon vital
capital, when interest alone should suffice ; it may be needed at
times to bridge a chasm, but habitual living beyond vital income
infallibly entails bankruptcy in health. The use of alcohol in
health seems practically restricted to purposes of sensuous gratifica-
tion on the part of those prepared to pay a round price for this
luxury. The three golden rules here are, — never drink before
breakfast, never drink alone, and never drink bad liquor ; their
observance may make even the abuse of alcohol tolerable. Serious
objections, for a naturalist at least, are that science, viewed through
a drinking-glass, seems distant and uncertain, while the pleasure of
drinking is immediate and unquestionable ; and that intemperance,
being an attempt to defy certain physical laws, is therefore
eminently unscientific.

SEC. V

REGISTRATION AND LABELLING

33

§ 5.— EEGISTEATION AND LABELLING

A mere Outline of a Field Naturalist’s Duties would be in-
excusably incomplete without mention of these important matters ;
and, because so much of the business of collecting must be left to be
acquired in the school of experience, I am the more anxious to give
explicit directions whenever, as in this instance, it is possible to do so.
Record your Observations Daily. — In one sense the specimens
themselves are your record, — prima facie evidence of your industry
and ability ; and if labelled as I shall presently advise, they tell no
small part of the whole story. But this is not enough ; indeed, I
am not sure that an ably conducted ornithological journal is not
the better half of your operations. Under your editorship of
labelling, specimens tell what they know about themselves j but
you can tell much more yourself. Let us look at a day’s
work : You have shot and skinned so many birds, and laid them
away labelled. You have made observations about them before
shooting, and have observed a number of birds that you did not
shoot. You have items of haunts and habits, abundance or scarcity;
of manners and actions under special circumstances, as of pairing,
nesting, laying, rearing young, feeding, migrating, and what not ;
various notes of birds are still ringing in your ears ; and finally,
you may have noted the absence of species you saw a while before,
or had expected to occur in your vicinity. Meteorological and topo-
graphical items, especially when travelling, are often of great assist-
ance in explaining the occurrences and actions of birds. Now you
know these things, but very likely no one else does ; and you know
them at the time, but you will not recollect a tithe of them in a few
weeks or months, to say nothing of years. Don’t trust your memory;
it will trip you up ; what is clear now will grow obscure ; what is
found will be lost. Write down everything while it is fresh in
your mind ; write it out in full ; time so spent will be time saved
in the end, when you offer your researches to the discriminating
public. Don’t be satisfied with a dry -as -dust item; clothe a
skeleton fact, and breathe life into it with thoughts that glow ; let
the paper smell of the woods. There’s a pulse in a new fact; catch
the rhythm before it dies. Keep off the quicksands of mere
memorandum — that means something “to be remembered,” which
is just what you cannot do. Shun abbreviations ; such keys rust
with disuse, and may fail in after times to unlock the secret that
should have been laid bare in the beginning. Use no signs intel-
ligible only to yourself ; your note-books may come to be overhauled
by others whom you would not wish to disappoint. Be sparing of
I)

34

FIELD ORNITHOLOGY

PART I

sentiment, a delicate thing, easily degraded to drivel ; crude enthusi-
asm always hacks instead of hewing. Beware of literary infelicities ;
“the written word remains,” it may he, after you have passed
away ; put down nothing for your friend’s blush, or your enemy’s
sneer ; write as if a stranger were looking over your shoulder.

Ornithological Book-keeping may he left to your discretion and
good taste in the details of execution. Each may consult his
preferences for rulings, headings, and blank forms of all sorts, as
well as particular modes of entry. But my experience has been
that the entries it is advisable to make are too multifarious to be
accommodated by the most ingenious formal ruling ; unless, indeed,
you make the conventional heading “ Kemarks ” disproportionately
wide, and commit to it everything not otherwise provided for. My
preference is decidedly for a plain page. I use a strongly bound
blank book, cap size, containing at least six or eight quires of good
smooth paper ; but smaller may be needed for travelling, even down
to a pocket note-book. I would not advise a multiplicity of books,
splitting up your record into different departments : let it be
journal and register of specimens combined. (The registry of you7'
own collecting has nothing to do with the register of your cabinet
of birds, which is sure to include a proportion of specimens from
other sources, received in exchange, donated, or purchased. I speak
of this beyond.) I have found it convenient to commence a day’s
record with a register of the specimens secured, each entry consisting
of a duplicate of the bird’s label (see beyond), accompanied by any
further remarks I have to make respecting the particular specimens ;
then to go on with the full of my day’s observations, as suggested
in the last paragraph. You thus have a register of collections in
chronological order, told off with an unbroken series of numbers,
checked with the routine label -items, and continually interspersed
with the balance of your ornithological studies. Since your private
field-number is sometimes an indispensable clew to the authentication
of a specimen after it has left your own hands, never duplicate it.
If 3mu are collecting other objects of natural history besides birds,
still have but one series of numbers ; duly enter your mammal, or
mineral, or whatever it is, in its place, with the number under which
it happens to fall. Be scrupulously accurate with these and all
other figures, as of dates and measurements. Always use black ink ;
lead-pencilling is never safe.

Labelling. — This should never be neglected. It is enough to
make a sensitive ornithologist shiver to see a specimen without that
indispensable appendage — a label. I am sorry to observe that the
routine labelling of most collections is far from being satisfactory.
A well-appointed label is something more than a slip of paper with
the bird’s name on it, and is still defective if, as is too often the

SEC. V

/LEGISLATION AND LABELLING

35

case, only the locality and collector are added. A complete label
records the following particulars : 1 . Title of the survey, voyage,
exploration, or other expedition (if any), during which the specimen
was collected. 2. Name of the person in charge of the same (and
it may be remarked that the less he really cares about birds, and
the less he actually interests himself to procure them, the more
particular he will be about this). 3. Title of the institution or
association (if any) under the auspices or patronage of which the
specimen was procured, or for which it is designed. 4. Name of
collector ; partly to give credit where it is due, but principally to
fix responsibility, and authenticate the rest of the items. 5.
Collectors number, referring to his note-book, as just explained; if
the specimen afterwards forms part of a general collection it
usually acquires another number by new registry ; the collector’s
then becoming the “original,” as distinguished from the “current,”
number. 6. Locality, perhaps the most important of all the items.
A specimen of unknown or even uncertain origin is worthless or
nearly so. Lamentable confusion has only too often arisen in
ornithological writings from vague or erroneous indications of
locality. I should say that a specimen not authentic in this
particular had better have its supposed origin erased. Nor will it
do to say simply, for instance, “North America” or “England.”
The general geographical distribution of birds being according to
recognised faunal areas, ornithologists generally know already the
quarter of the globe from which any bird comes ; the locality of
particular specimens, therefore, should be fixed down to the very
spot. If this be obscure, add the name of the nearest place to be
found on a fairly good map, giving distance and direction. 7. Date
of collection, — day of the month, and year. Among other reasons for
this may be mentioned the fact that it is often important to know
what season a particular plumage indicates. 8. Sex, and if possible
also age, of the specimen, — an item that bespeaks its own import-
ance. Ornithologists of all countries are agreed upon certain signs
to indicate the sex. These are ; f for male, $ for female, — the
symbols respectively of Mars and Venus. Immaturity is often
denoted by the sign ^ ; thus, f young male. Or, we may write
? ad., $ yg., for adult female, young female, respectively. It is
preferable, however, to use the language of science, not our
vernacular, and say f juv. (juvenis, young). Nupt. signifies breed-
ing plumage ; hornot. means a bird of the year. 9. Measure-
ments of length, and of extent of wings ; the former can only be
obtained approximately, and the latter not at all, from a prepared
specimen. 10. Colour of the eyes, and of the bill, feet, or other
naked or soft parts, the tints of which may change in drying.
11. Miscellaneous particulars, such as contents of stomach, special

36

FIELD ORNITHOLOGY

PART I

circumstances of capture, vernacular name, etc. 12. Scientific name
of the bird. This is really the least important item of all, though
generally thought to take precedence. But a bird labels itself, so
to speak ; and nature’s label may be deciphered at any time. In
fact, I would enjoin upon the collector not to write out the supposed
name of the bird in the field, unless the species is so well known as
to be absolutely unquestionable. Proper identification, in any case
to which the slightest doubt may attach, can only be made after
critical study in the closet with ample facilities for examination
and comparison. But it is always well to note on the label the
local vernacular name ; for native names, especially un-English
ones, may become valuable items of information. The first eight
items above, and the twelfth, usually constitute the face or
obverse of a label ; the rest are commonly written on the back or
reverse side. Labels should be of light cardboard, or very stiff
writing paper ; they may be dressed attractively, as fancy
suggests j the general items of a large number of specimens are best
printed ; the special ones must of course be written. Shape is
immaterial. A slip about three inches long and two-thirds of
an inch wide will do very well for anything, from a hawk to
a humming-bird. Something like the shipping -tag used by
merchants is excellent, particularly for larger objects. It seems
most natural to attach the string to the left-hand end. The slip
should be tied so as to swing just clear of the bird’s legs, but not
loose enough to dangle several inches, for in that case the labels
are continually tangling with each other when the birds are laid
away in drawers. The following forms show the face and back of
the last label I happened to write before these lines were originally
penned; they represent the size and shape that I find most
convenient for general purposes ; while the legend illustrates every
one of the twelve items above specified.

i Explorations in Dakota. Dr. Elliott Cones, U.S.A.

‘3

1 No. 2655. Buteo borealis (Gm.) V. ? juv.

'S Fort Randall, Missouri River.

OQ

Oct. 29, 1872.

o’

Obverse.

23.00 X 53.00 X 17.50. — Eyes yellowish - gray ;
darker at tip ; cere wax - yellow ; tarsi dull
bluish - black. Stomach contained portions of
large tapeworm.

bill horn - blue,
yellowish ; claws
a rabbit ; also, a

Reverse.

SEC. V

REGISTRATION AND LABELLING

37

Directions for Measurement may be inserted here, as this
matter pertains rightfully to the recording of specimens. The
following instructions apply not only to length and extent, but to
the principal other dimensions, which may be taken at any time.
For large birds, a tape-line showing inches and fourths will do ; for
smaller ones, a foot-rule graduated for inches and eighths, or better,
decimals to hundredths, must be used ; and for all nice measurements
the dividers are indispensable. Length : Distance between the tip
of the bill and end of the longest tail-feather. Lay the bird on its
back on the ruler on a table ; take hold of the bill with one hand
and of both legs with the other ; pull with reasonable force to get
the curve all out of the neck ; hold the bird thus with the tip of
the bill flush with one end of the ruler, and see where the end of
the tail points. Put the tape-line in place of the ruler, in the same
way, for larger birds. Extent: Distance between the tips of the
outspread wings. They must be fully outstretched, with the bird
on its back, crosswise on the ruler, its bill pointing to your
breast. Take hold of right and left metacarpus with the thumb and
forefinger of your left and right hand respectively, stretch with
reasonable force, getting one wing -tip flush with one end of the
ruler, and see how far the other wing -tip reaches. With large
birds pull as hard as you please, and use the table, floor, or side of
the room ; mark the points and apply tape-line. Length of wing :
Distance from the carpal angle formed at the bend of the wing to
the end of the longest primary. Take it with compasses for small
birds. In birds with a convex wing, do not lay the tape-line over
the curve, but under the wing in a straight line. This measure-
ment is the one called, for short, “the wing.” Length of tail:
Distance from the roots of the rectrices to the end of the longest one.
Feel for the pope’s-nose ; in either a fresh or dried specimen there is
more or less of a palpable lump into which the tail-feathers stick.
Guess as near as you can to the middle of this lump ; place the end
of the ruler opposite this point, and see where the tip of the longest
tail-feather comes. Length of hill: Some take the curve of the
upper mandible ; others the side of the upper mandible from the
feathers; others the gape, etc. I take the chord of the culmen.
Place one foot of the dividers on the culmen just where the feathers
end ; no matter whether the culmen runs up on the forehead, or
the frontal feathers run out on the culmen, and no matter whether
the culmen is straight or curved. Then with me the length of
the hill is the shortest distance from the point just indicated to the
tip of the upper mandible : measure it with the dividers. In a
straight bill of course it is the length of the culmen itself ; in a
curved bill, however, it is quite another thing. Length of tarsus:
Distance between the joint of the tarsus with the leg above, and

38

FIELD ORNITHOLOGY

PART I

that with the first phalanx of the middle toe below. Measure it
always with dividers, and in front of the leg. Length of toes :
Distance in a straight line along the upper surface of a toe from the
point last indicated to the root of the claw on top. Length of
toe is to be taken without the claw, unless otherwise specified.
Length of the claws: Distance in a straight line from the point
last indicated to the tip of the claw. Length of head is often a
convenient dimension for comparison with the bill. Set one foot of
the dividers over the base of the culmen (determined as above)
and allow the other to slip snugly down over the arch of the
occiput.

§ 6.— INSTEUMENTS, MATEEIALS, AND FIXTUEES FOE
PEEPAEIND BIEDSKINS

Instruments. — The only indispensable instrument is a pair of
scissors or a knife ; practically, you want both of these, a pair of
spring-forceps, and a knitting-needle, or some similar wooden or
ivory object. I have made hundreds of birdskins consecutively
without touching another tool. Persicos odi, 'puer^ apparatus I I
always mistrust the emphasis of a collector who makes a flourish of
instruments. You might be surprised to see what a meagre, shabby-
looking kit our best taxidermists work with. Stick to your scissors,
knife, forceps, and needle. But you may as well buy, at the outset,
a common dissecting-case, such as medical students begin business
with j it is very cheap, and if there are some unnecessary things in
it, it makes a nice little box in which to keep your tools. The case
contains, among other things, several scalpels, just the knives you
want ; a “ cartilage-knife,” which is nothing but a stout scalpel, suit-
able for large birds ; the best kind of scissors for your purpose, with
short blades and long handles — if kneed at the hinge so much
the better ; spring forceps, the very thing ; a blow-pipe, useful in
many ways and answering instead of a knitting-needle ; and some
little steel hooks, chained together, which you may want to use.
But you will also require, for large birds, a very heavy pair of scissors,
or small shears, short-bladed and long-handled, and a stout pair of
bone nippers. Have some pins and needles ; surgical needles, which
cut instead of punching, are the best. Get a hone or strop, if you
wish, and a feather-duster. Use of scissors requires no comment,
and I would urge their habitual employ instead of the knife-blade ;
I do nine-tenths of my cutting with scissors, and find it much the
easiest. A double-lever is twice as effective as a single one. More-
over, scalpels need constant sharpening ; mine are generally too dull

SEC. VI INSTRUMENTS, ETC., FOR PREPARING BIRD SKINS 39

to cut much with, and I suppose I am like other people — while
scissors stay sharp enough. The flat, thin ivory or ebony handle of
the scalpel is about as useful as the blade. Finger-nails, which were
made before scalpels, are a mighty help. Forceps are almost indis-
pensable for seizing and holding parts too small or too remote to be
grasped by the fingers. The knitting-needle is wanted for a specific
purpose noted beyond. The shears or nippers are only needed for
what the ordinary scissors are too weak to do.

Materials. — {a) For stuffing. “What do you stuff ’em with?”
is usually the first question of idle curiosity about taxidermy, as if
that were the great point j whereas the stuffing is so small a matter
that one might reply, “ Anything, except brickbats ! ” But if stuffing
birds were the final cause of cotton, that admirable substance could
not be more perfectly adapted than it is to the purpose. Ordinary
raw cotton-batting or wadding is what you want. When I can get
it I never think of using anything else for small birds. I would use
it for all birds were expense no object. Here tow comes in ; there
is a fine, clean, bleached article of tow prepared for surgical dressings ;
this is the best, but any will do. Some say chop your tow fine ; this
is harmless, but unnecessary. A crumpled newspaper, wrapped with
tow, is first-rate for a large bird. Failing cotton or tow, any soft,
light, dry, vegetaUe substance may be made to answer, — rags, paper,
crumbled leaves, fine dried grass, soft fibrous inner bark, etc. ; the
down of certain plants, as thistle and silkweed, makes an exquisite
filling for small birds. But I will qualify my remark about brick-
bats by saying : Never put hair, wool, feathers, or any other animal
substance in a birdskin ; far better leave it empty : for, as we shall see
in the sequel, bugs come fast enough, without being invited into a
snug nest. if) For preserving. Arsenic, — not the pure metal
properly so called, but arsenic of the shops, or arsenious acid, — is
the great preservative. Use dry powdered arsenic, plenty of it, and
nothing else. There is no substitute for arsenic worthy of the name,
and no preparation of arsenic so good as the simple substance.
Various kinds of “arsenical soap” were and may still be in vogue;
it is a nasty, greasy substance ; and although efiicacious enough, there
is a very serious hygienic objection to its use.^ Arsenic, I need not
say, is a violent irritant poison, and must therefore be duly guarded,
but may be used with perfect impunity. It is a very heavy substance,
not appreciably volatile at ordinary temperatures, and therefore not

^ “ Strange as it may appear to some, I would say avoid especially all the so-called
arsenical soaps ; they are at best but filthy preparations ; besides, it is a fact to which
I can bear painful testimony that they are, especially when applied to a greasy skin,
poisonous in the extreme. I have been so badly poisoned, while working upon the
skins of some fat water birds that had been prepared with arsenical soap, as to be
made seriously ill, the poison having worked into the system through some small
wounds or scratches on my hand. Had j)ure arsenic been used in preparing the skins,

40

FIELD ORNITHOLOGY

PART I

liable, as some suppose, to be breathed, to any perceptible, much
less injurious, extent. It will not at once enter the pores of healthy
unbroken skin ; so it is no matter if it gets on the fingers. The
exceedingly minute quantity that may be supposed to find its way
into the system in the course of time is believed by many competent
physicians to be rather beneficial as a tonic. I will not commit
myself to this ; for, though I have never felt better than when
working daily with arsenic, I do not know how much my health
was improved by the outdoor exercise always taken at the same
time. The simple precautions are, not to let it lie too long in con-
tact with the skin, nor get into an abrasion, nor under the nails.
It will convert a scratch or cut into a festering sore of some little
severity ; while if lodged under the nails it soon shows itself by
soreness, increased by pressure ; a white speck appears, then a tiny
abscess forms, discharges, and gets well in a few days. Your pre-
cautions really respect other persons more than yourself ; the
receptacle should be conspicuously labelled “ POISON ! ” Arsenic
is a good friend ; besides preserving our birds, it keeps busybodies
and meddlesome folks away from the scene of operations, by raising
a wholesome suspicion of the taxidermist’s surroundings. It may
be kept in the tin pots in which it is usually sold ; but some shallower,
broader receptacle is more convenient. A little drawer say 6x6
inches, and an inch deep, to slip under the edge of the table, or a
similar compartment in a large drawer, will be found handy. A
salt-spoon, or little wooden shovel whittled like one, is nice to use it
with, though it is in fact generally taken up with the handle of the
scalpel. As stated, there is no substitute for arsenic ; but at a pinch
you can make temporary shift with the following, among other
articles : table salt, or saltpetre, or charcoal strewn plentifully ;
strong solution of corrosive sublimate, brushed over the skin inside ;
creosote \ impure carbolic acid — these last two are quite efficacious,
but they smell horribly for an indefinite period. A bird threatening
to decompose before you are ready to skin it, may be saved for a
while by injecting weak carbolic acid or creosote down the throat
and up the fundament ; or by disembowelling, and filling the cavity
with powdered charcoal, (c) For cleansing. Gypsum is an almost
indispensable material for cleansing soiled plumage. Gypsum is
properly native hydrated sulphate of lime ; the article referred to
is “plaster of Paris” or gypsum heated up to 260° F. (by which the
water of crystallisation is driven off) and then finely pulverised.

the effect would not have been as had, although grease and arsenic are generally a
blood-poison in sovie degree ; but when combined with ‘ soap ’ the effect, at least as
far as my experience goes, is much more injurious” (Maynard, Guide, p. 12). In
indorsing this, I would add that the combination is the more poisonous, in all prob-
ability, simply because the soap, being detersive, mechanically facilitates the entrance
of the poison, without, however, chemically increasing its virulence.

SEC. VI INSTRUMENTS, ETC., FOR PREPARING BIRDS KINS 41

When mixed with water it soon solidifies, the original hydrate being
again formed. The mode of using it is indicated beyond. It is
most conveniently kept in a shallow tray, say a foot square, and an
inch or two deep, which had better, furthermore, slide under the
table as a drawer ; or form a compartment of a larger drawer.
Keep gypsum and arsenic in different-looking receptacles, not so much
to keep from poisoning yourself, as to keep from not poisoning a
birdskin. They look much alike, and skinning becomes such a
mechanical process that you may get hold of the wrong article when
your thoughts are wandering in the woods. Gypsum, like arsenic,
has no worthy rival in its own field ; some substitutes, in the order
of their applicability, are : corn-meal, probably the best thing after
gypsum ; calcined magnesia (very good but too light — it floats in
the air, and makes you cough) ; bicarbonate of magnesia ; powdered
chalk prepared chalk,” creta prceparata of the drug shops, is the best
kind) ; fine wood-ashes ; clean dry loam. No article, however
powdery when dry, that contains a glutinous principle, as for instance
gum-arabic or flour, is admissible, {d) For wrapping, you want a
thin, pliable, strong paper toilet-paper is the very best ; newspaper
is pretty good. For making the cones or cylinders in which bird-
skins may be set to dry, a stiffer article is required ; writing paper
answers perfectly.

Naturalists habitually carry a Pocket Lens, much as other
people do a watch. You will find a magnifying glass very con-
venient in your search for the sexual organs of small birds when
obscure, as they frequently are, out of the breeding-season ; in
picking lice from plumage, to send to your entomological friend,
who will very likely pronounce them to be of a new species ; and for
other purposes.

Fixtures. — When travelling, your fixtures must ordinarily be
limited to a collecting-chest ; you will have to skin birds on the top
of this, on the tail-board of a wagon, or on your lap, as the case may
be. The chest should be very substantial — iron-bound is best ;
strong as to hinges and lock— and have handles. A good size is
30x18x18 inches. Let it be fitted with a set of trays; the
bottom one say four inches deep ; the rest shallower ; the top one
very shallow, and divided into compartments for your tools and
materials, unless you fix these on the under side of the lid. Start
out with all the trays full of cotton or tow. At home have a room
to yourself, if possible ; taxidermy makes a mess to which your wife
may object, and arsenic must not come in the way of children. At
any rate have your own table. Great cleanliness is indispensable,
especially when doing much work in hot weather, for the j^lace soon
smells sour if neglected. I use no special receptacle for offal, for
this only makes another article to be cleaned ; lay down a piece of

42

FIELD ORNITHOLOGY

PART I

paper for the refuse, and throw the whole away. A perfectly smooth
surface is desirable. I generally have a large pane of window-glass
on the table before me. It will really be found advantageous to
have a scale of inches scratched on the edge of the table ; only a
small part of it need be fractionally subdivided ; this replaces the
foot-rule and tape-line, just as the tacks of a dry-goods counter answer
for the yard-stick. You will find it worth while to rig some sort of a
derrick arrangement, w^hich you can readily devise, on one end of
the table, to hitch your hook to, if you hang your birds up to skin
them j they should swing clear of everything. The table should
have a large general drawer, with a little drawer for gypsum and
arsenic, unless these be kept elsewhere. Stuffing may be kept in a
box under the table, and make a nice footstool ; or in a bag slung
to the table leg.

§ 7.— HOW TO MAKE A BIRDSKIN

{a) The Regular Process

Lay the Bird on its Back, the bill pointing to your right ^
elbow. Take the scalpel like a pen, with edge of blade uppermost,
and run a straight furrow through the feathers along the middle
line of the belly, from end of the breast -bone to the vent. Part
the feathers completely, and keep them parted.^ Observe a strip
of skin either perfectly naked, or only covered with short down ;
this is the line for incision. Take scissors, stick in the pointed
blade just over the end of the breast-bone, cut in a straight line
thence to and into the vent ; cut extremely shallow. ^

Take the forceps in your left hand, and scalpel in your right,
both held pen-wise, and with the forceps seize and lift up one of
the edges of the cut skin, gently pressing away the belly-walls with

^ Reverse this and following directions for position, if you are left-handed.

^ The motion is exactly like stroking the right and left sides of a moustache
apart ; you would never dress the hairs smoothly away from the middle line, by
poking from ends to root ; nor will the feathers stay aside, unless stroked away from
base to tip.

^ The skin over the belly is thin as tissue paper in a small bird ; the chances are
you will at first cut the walls of the belly too, opening the cavity ; this is no great
matter, for a pledget of cotton will keep the bowels in ; nevertheless, try to divide
skin only. Reason I'or cutting into vent : this orifice makes a nice natural termina-
tion of the incision, buttonhole-wise, and may keep the end of the cut from tearing
around the root of the tail. Reason for beginning to cut over the edge of the breast-
bone : the muscular walls of the belly are very thin, and stick so close to the skin
that you may be in danger of attemiDting to remove them with the skin, instead of
removing the skin from them ; whereas you cannot remove anything but skin from
over the breast-bone, so you have a guide at the start. You can tell skin from
belly-wall, by its livid, translucent whitishness instead of redness.

SEC. VII

HOW TO MAKE A BIRD SKIN

43

the scalpel-point; no cutting is required; the skin may be peeled
off without trouble. Skin away till you meet an obstacle ; it is
the thigh. Lay down the instruments ; with your left hand take
hold of the leg outside at the shank ; put your right forefinger
under the raised fiap of skin, and feel a bump ; it is the knee; push
up the leg till this bump comes into view ; hold it so. Take the
scissors in your right hand ; tuck one blade under the concavity of
the knee, and sever the joint at a stroke ; then the thigh is left
with the rest of the body, while the rest of the leg is dissevered and
hangs only by skin. Push the leg farther up till it has slipped
out of its sheath of skin, like a finger out of a glove, down to the
heel-joint. You have now to clear off the flesh and leave the bone
there ; you may scrape till this is done, but there is a better way.
Stick the closed points of the scissors in among the muscles just
below the head of the bone, then separate the blades just wide
enough to grasp the bone ; snip off its head ; draw the head to one
side ; all the muscles follow, being there attached ; strip them
downward from the bone ; the bone is left naked, with the muscle
hanging by a bundle of tendons (“leaders”) at its foot ; sever these
tendons collectively at a stroke. This whole performance will
occupy about three seconds, after practice ; and you may soon
discover you can nick off the head of the bone of a small bird with
the thumb-nail. Draw the leg-bone back into its sheath, and leave
it. Eepeat the foregoing steps on the other side of the bird. If
you are bothered by the skin-flaps settling against the belly-walls,
insert a fluff of cotton. Keep the feathers out of the wound ; cotton
and the moustache movement will do it. Next you must sever the
tail from the body, leaving a small “ pope’s-nose ” for the feathers
to stay stuck into. Put the bird in the hollow of your lightly
closed left hand, tail upward, belly toward you ; or, if too large for
this, stand it on its breast on the table in similar position. Throw
your left forefinger across the front (under side) of the tail, pressing
a little backward ; take the scissors, cut the end of the lower bowel
free first, then peck away at bone and muscle with cautious snips,
till the tail-stump is dissevered from the rump, and the tail hangs
only by skin. You will soon learn to do it all at one stroke ; but
you cannot be too careful at first ; you are cutting right down on
to the skin over the top of the pope’s-nose, and if you divide this,
the bird will part company with its tail altogether. Now you have
the rump-stump protruding naked; the legs dangling on either
side ; the tail hanging loose down over the bird’s back. Lay down
scissors, take up forceps ^ in your left hand ; with them seize and

^ Or at this stage you may instead stick a hook into a firm part of the rump,
and hang up the bird about the level of your breast ; you thus have both hands
free to work with. This is advisable with all birds too large to be readily taken in

44

FIELD ORNITHOLOGY

PART I

hold the stump of the rump ; and with point or handle of scalpel
in the other hand, with finger-tips, or with thumb-nail (best), gently
press down on and peel away skind No cutting will be required
(usually) till you come to the wings : the skin peels off (usually) as
easily as an orange-rind ; as fast as it is loosened, evert it ; that is,
make it continually turn itself more and more completely inside
out. Work thus till you are stopped by the obtruding wings.^
You have to sever the wing from the body at the shoulder, just as
you did the leg at the knee, and leave it hanging by skin alone.
Take your scissors,^ as soon as the upper arm is exposed, and cut
through flesh and bone alike at one stroke, a little below (outside
of) the shoulder -joint. Do the same with the other wing. As
soon as the wings are severed the body has been skinned to the
root of the neck ; the process becomes very easy ; the neck almost
slips out of its sheath of itself ; and if you have properly attended
to keeping the feathers out of the wound and to continual eversion
of the skin, you now find you have a naked body connected dumb-
bell-wise by a naked neck to a cap of reversed skin into which
the head has disappeared, from the inside of which the legs and
wings dangle, and around the edges of which is a row of plumage
and a tail.^ Here comes up an important consideration : the skin,
plumage, legs, wings, and tail together weigh something, — enough
to stretch ^ unduly the skin of the neck, from the small cylinder of

hand, and will help you, at first, with any bird. But there is really no use of it
with a small bird, and you may as well learn the best way of working at first as
afterward.

^ The idea of the whole movement is exactly like ungloving your hand from the
wrist, by turning the glove inside out to the very finger tips. Some say, pull
off the skin ; I say never pull a bird’s skin under any circumstances : push it off,
always operating at lines of contact of skin with body, never upon areas of skin
already detached.

^ The elbows will get in your, way before you reach the point of attack, namely,
the shoulder, unless the wings were completely relaxed (as was essential, indeed, if
you measured alar expanse correctly). Think what a difference it would make, were
you skinning a man through a slit in the belly, whether his arms were stretched
above his head or pinned against his ribs. It is just the same with a bird. When
properly relaxed the wings are readily pressed away toward the bird’s head, so that
the shoulders are encountered before the elbows.

^ Shears will be required to crash through a large arm-bone. Or, you may with
the scalpel unjoint the shoulder. The joint will be found higher up and deeper
among the breast muscles than you might suppose, unless you are used to carving
fowls at table. With a small bird, you may snap the bone with the thumb-nail and
tear asunder the muscles in an instant.

You find that the little straight cut you made along the belly has somehow
become a hole larger than the greatest girth of the bird ; be undismayed ; it is all
right.

^ If you have up to this point properly pushed off the skin instead of pulling it,
there is as yet probably no stretching of any consequence ; but, in skinning the
head, which comes next, it is almost impossible for a beginner to avoid stretching to
an extent involving great damage to the good looks of a skin. Try your utmost, by
delicacy of manipulation at the lines of contact of skin with flesh, and only there, to

SEC. VII

HOW TO MAKE A BIRDS KIN

45

which they are now suspended ; the whole mass must be supported.
For small birds, gather it in the hollow of your left hand, letting the
body swing over the back of your hand out of the way ; for large
ones, rest the affair on the table or your lap. To skin the head,
secure the body in the position just indicated, by confining the neck
between your left thumb and forefinger ; bring the right fingers
and thumb to a cone over the head, and draw it out with gentle
force ; or, holding the head itself between the left thumb and
forefinger, insert the handle of the scalpel between the skin and
skull, and pry a little, to enlarge the neck-cylinder of skin enough
to let the head pass. It will generally^ slip out of its hood very
readily, as far as its greatest diameter ; ^ there it sticks, being in
fact pinned by the ears. Still holding the bird as before, with the
point of the scalpel handled like a nut-picker, or with your thumb-
nail, detach the delicate membrane that lines the ear-opening ; do
the same for the other ear. The skull is then shelled out to the
eyes, and will skin no farther of its own accord, being again
attached by a membrane, around the border of the eye-socket.
Holding the scalpel as before, run its edge around an arc (a semi-
circle is enough to let you into the orbit) of the circumference,
dissevering the membrane from the bone. Eeverse the scalpel, and
scoop out the eyeball with the end of the handle ; you bring out
the eye betwixt the ball of your thumb and the handle of the
instrument, tearing apart the o]Dtic nerve and the conjunctival
tissue, but taking care not to open the eyeball^ or lacerate the
eyelids. Do the same with the other eye. The head is then
skinned far enough ; there is no use of getting quite to the base of
the bill. You have now to get rid of the brain and fiesh of the
nape and jaws,^ and leave most of the skull in ; the cranial dome
makes the only perfect “ stuffing ” for the skin of the head. This
is all done at once by only four particular cuts. Hold the head

prevent lengthwise stretching. Crosswise distension is of no consequence ; in fact
more or less of it is usually required to skin the head, and it tends to counteract the
ill effect of undue elongation.

^ The special case of head too large for the calibre of the neck is treated
beyond.

^ And you will at once find a great apparent increase of amount of free skin in
your hand, owing to release and extension of all that was before shortened in length
by circular distension, in enlargement of the neck-cylinder.

^ An eyeball is much larger than it looks from the outside ; if you stick the
instrument straight into the socket, you may punch a hole in the ball and let out
the water — a very disagreeable complication. Insinuate the knife-handle close to the
rim of the socket, and hug the wall of the cavity throughout.

You may of course at this stage cut off the neck at the nape, punch a hole in
the base of the skull, dig out the brains, and scrape away at the jaw-muscles till you
are satisfied or tired ; an unnecessary job, during which the skin may have become
dry and shrivelled and hard to turn right side out. The operation described in the
text may require five seconds, perhaps.

46

FIELD ORNITHOLOGY

PART I

between your left thumb and fingers, the bill pointing towards you,
the bird’s palate facing you ; you observe a space bounded behind
by the base of the skull where the neck joins, in front by the floor
of the mouth, on either side by the prongs of the under jaw, — these
last especially prominent. Take the scissors ; stick one blade just
inside one branch of the lower jaw, thence into the eye-socket
which lies below (the head being upside down), thence into the
brain -box; make a cut parallel with the jaw, just inside of it,
bringing the upper scissor-blade perpendicularly downward, crash-
ing through the skull just inside of the angle of the jaw. Duplicate
this cut on the other side. Connect the anterior ends of these cuts
by a transverse one across the floor and roof of the mouth. Connect
the posterior ends of the side cuts by one across the back of the skull
near its base, — ^just where the nape-muscle ceases to override the
cranium. You have enclosed and cut out a squarish-shaped mass of
bone and muscle, and, on gently pulling the neck (to which of course
it remains attached) the whole affair comes out, bringing the brain
with it, but leaving the entire roof of the skull supported on a scaffold-
ing of jaw-bone. It only remains to skin the wings. Seize the arm-
stump with fingers or forceps ; the upper arm is readily drawn from
its sheath as far as the elbow ; but the wing must be skinned to
the wrist (carpus — “ bend of the wing ”) ; yet it will not come out
easily, because the secondary quills grow to one of the forearm
bones (the ulna), pinning down the skin the whole way along a
series of points. To break up these connections, hold the upper
arm firmly with the left thumb and forefinger, the convexity of the
elbow looking towards you; press the right thumb-nail closely
against the back edge of the ulna, and strip downward, scraping
the bone with the nail the whole way. If you only hit the line of
adhesions, there is no trouble at all about this. Now you want to
leave in one of the two forearm bones, to preserve sufficiently the
shape of the limb, but to remove the other, with the upper-arm
bone and all the flesh. It is done in a moment : stick the point of
the scissors between the heads of the two forearm bones, and "cut
the hinder one (ulna) away from the elbow ; then the other fore-
arm bone (radius), bearing on its near end the elbow and the whole
upper-arm, is to be stripped away from the ulna, taking with it the
flesh of the forearm, and to be cut off at its far end close to the
wrist-joint, one stroke severing the bone and all the tendons that
pass over the wrist to the hand ; then the ulna, bare of flesh, is
alone left in, attached at the wrist. Draw gently on the wing from
the outside till it slips into the natural position whence you everted
it. Do the same for the other wing. This finishes the skinning
process. The skin is now to be turned right side out. Begin any
way you please, till you see the point of the bill reappearing among

SEC. VII

HOW TO MAKE A BIRD SKIN

47

the feathers ; seize it with fingers or forceps, as convenient, and use
it for gentle traction. But by no means pull it out by holding on
to the rear end of the skin — that would infallibly stretch the skin.
Holding the bill, make a cylinder of your left hand and coax the
skin backward with a sort of milking motion. It will come easily
enough, until the final stage of getting the head back into its skull-
cap ; this may require some little dexterity ; but you cannot fail to
get the head in, if you remember what you did to get it out. When
this is fairly accomplished, you for the first time have the pleasure
of seeing something that looks like a birdskin. Your next care is
to apply arsenic. Lay the skin on its back, the opening toward
you and wide spread, so the interior is in view. Bun the scalpel-
handle into the neck to dilate that cylinder until you can see the
skull ; find your way to the orifices of the legs and wings ; expose
the pope’s-nose ; thus you have not only the general skin surface,
but all the points where some traces of flesh were left, fairly in
view. Put in arsenic ; send some down the neck, making sure it
reaches and plentifully besprinkles the whole skull ; drop a little
in each wing-hole and leg-hole ; leave a small pile at the root of the
tail ; strew some more over the skin at large. The simple rule is,
put in as much arsenic as will stick anywhere. Then close the
opening, and shake up the skin ; move the head about by the bill ;
rustle the wings and move the legs ; this distributes the poison
thoroughly. If you have got in more than is necessary, as you may
judge by seeing it piled up dry, anywhere, hold the skin with the
opening downward over the poison-drawer, and give it a flip and let
the superfluous powder fall out. Now for the “make-up,” upon
which the beauty of the preparation depends. First get the empty
skin into good shape. Let it lie on its back ; draw it straight out
to its natural length. See that the skin of the head fits snugly ;
that the eyes, ears, and jaws are in j^lace. Expand the wings to
make sure that the bone is in place, and fold them so that the
quills override each other naturally ; set the tail-feathers shingle-
wise also ; draw down the legs and leave them straddling wide
apart. Give the plumage a preliminary dressing ; if the skin is free
from kinks and creases, the feathers come naturally into place ;
particular ones that may be awry should be set right, as may be
generally done by stroking, or by lifting them free repeatedly, and
letting them fall ; if any (through carelessness) remain turned into
the opening, they should be carefully picked out. Eemove all traces
of gypsum or arsenic with the feather-duster. The stuffing is to be
put in through the opening in the belly ; the art is to get in just
enough, in the right places. It would never do to push in pellets
of cotton, as you would stuff a pillow-case, till the skin is filled up ;
no subsequent skill in setting could remove the distortion that

48

FIELD ORNITHOLOGY

PART I

would result.! It takes just four pieces of stuffing — one for each
eye, one for the neck, and one for the body ; while it requires
rather less than half as much stuffing as an inexperienced person
might suppose. Take a shred of cotton that will make a tight ball
as large as the bird’s eye ; stick it on the end of your knitting-
needle, and by twirling the needle whilst the cotton is confined in
your finger tips, you make a neat ball. Introduce this through the
belly-opening into the eye-socket; if you have cut away skull
enough, as already directed, it will go right in ; disengage the
needle with a reverse twirl, and withdraw it. Take hold of the
bill with one hand, and with the forceps in the other, dress the eye-
lids neatly and naturally over the elastic substance within. Eepeat
for the other eye. Take next a shred of cotton that will roll into
a firm cylinder rather less than the size of the bird’s neck. Eoll it
on the needle much as you did the eyeball, introduce it in the same
way, and ram it firmly into the base of the skull ; disengage the
needle by twirling it the other way, and withdraw it, taking care
not to dislodge the cotton neck. If now you peep into the skin
you will see the end of this artificial neck ; push it up against the
skin of the breast, — it must not lie down on the back between the
shoulders.^ The body- wad comes next, to imitate the size and shape
of the bird’s trunk. Take a mass of cotton you think will be
enough, and take about half of this ; that will be plenty (cotton is
very elastic). It should make a tolerably firm ball, rather egg-
shaped, swelling at the breast, smaller behind. If you simply
squeeze up the cotton, it will not stay compressed ; it requires a
motion something like that which bakers employ to knead dough
into the shape of a loaf. Keep tucking over the borders of the
cotton till the desired shape and firmness are attained. Insert the

1 For any ordinary bird up to tlie size of a crow, it is often directed that the
leg-bones and wing-bones be wrapped with cotton or tow. I should not think of
putting anything around the wing-bones of any bird up to the size of an eagle, swan,
or pelican. Examination of a skinned wing will show how extremely compact it is,
except just at the shoulder. What you remove will never make any difference from
the outside, while you would almost inevitably get in too much, not of the right
shape, and make an awkward bulging no art would remedy ; I say, then, leave the
wings of all but the largest birds emjpty, and put in very little cotton under any cir-
cumstances. As for legs, the whole host of small perching birds need no wrapping
whatever ; depend upon it you will make a nicer skin without wrapping. But large
birds and those with very muscular or otherwise prominent legs must have the
removal of flesh compensated. I treat of these cases beyond.

2 Although a bird’s neck is really, of course, in direct continuation of the back-
bone, yet the natural sigmoid curve of the neck is such that it virtually takes depart-
ure rather from the breast, its lower curve being received between the prongs of the
merrythought. This is what we must imitate instead of the true anatomy. If you
let the end of the neck lie between the shoulders, it will infallibly press them apart,
so that the interscapular plumage cannot shingle over the scapular feathers as it
should, and a gaping place, showing down or even naked skin, will result. Likewise,
if the neck be made too large (the chances are that way at first), the same result
follows. These seemingly trifling points are very important.

SEC. VII

HO FT TO MAKE A BIRD SKIN

49

ball between the blades of the forceps in such way that the instru-
ment confines the folded-over edges, and with a wriggling motion
insinuate it aright into the body. Before relaxing the forceps, put
your thumb and forefinger in the bird’s armpits, and pinch the
shoulders together till they almost touch \ this is to make sure that
there is no stuffing between the shoulders, — the whole mass lying
breastwards. Loosen the forceps and withdraw them. If the ball
is rightly made and tucked in, the elasticity of the cotton will
chiefly expend itself in puflSng out the breast, which is just what is
wanted. Be careful not to push the body too far in ; if it impacts
against the skin of the neck, this will infallibly stretch, driving the
shoulders apart, and no art will remedy the unsightly gap resulting.
You see I dwell on this matter of the shoulders ; the whole knack
of stuffing correctly focuses just over the shoulders. If you find you
have made the body too large, pull it out and make a smaller one ;
if it fits nicely about the shoulders, but is too long to go in, or too
puffy over the belly, let it stay, and pick away shreds at the open
end till the redundancy is remedied. Your bird is now stuffed.
Close the opening by bringing the edges of the original cut together.
There is no use of sewing up the cut for a small bird ; if the
stufling is correct, the feathers will hide the opening ; and if they
do not, it is no matter. You are not making an object for a show-
case, but for a naturalist’s cabinet. Supposing you to have been so
far successful, little remains to be done ; the skin already looks very
much like a dead bird j you have only to give the finishing touches,
and “ set ” it. Fixing the wings nicely is a great point. Fold each
wing closely; see that the carpal bend is well defined, that the
coverts show their several oblique rows perfectly, that all the quills
override each other like shingles. Tuck the folded wings close up
to the body — rather on the bird’s back than along its sides ; see
that the wing tips meet over the tail (under the tail as the bird lies
on its back) ; let the carpal angle nestle in the plumage ; have the
shoulders close together, so that the interscapular feathers shingle
over the scapulars. If the wing be pressed in too tightly, the
scapulars will rise up on end ; there must be neither furrow nor
ridge about the insertion of the wings ; everything must lie perfectly
smooth. At this stage of the process lift up the skin gingerly, and
let it slip head first through one hand after the other, pressing here
or there to correct a deformity, or uniformly to make the whole
skin compact. The wings set, next bring the legs together, so that
the bones within the skin lie parallel with each other ; bend the
heel -joint a little, to let the tarsi cross each other about their
middle ; lay them sidewise on the tail, so that the naturally flexed
toes lie flat, all the claws facing each other. See that the neck is
perfectly straight, and, if anything, shortened rather than out-
E

50

FIELD ORNITHOLOGY

PART I

stretched ; have the crown of the head flat on the table, the bill
pointing straight forward/ the mandibles shut tightly.^ Never
attempt any fancy attitudes with a birdskin ; the simpler and
more compactly it is made up the better.^ Finally, I say, hang
over your bird (if you have time) ; dress better the feathers that
were well dressed before ; perfect every curve ; finish caressingly,
and put it away tenderly, as you hope to be shriven yourself when
the time comes.

There are several ways of laying a birdskin. A common, easy,
and slovenly way is to thrust it head first into a paper cone ; but
it makes a hollow-chested, pot-bellied object, unpleasant to see, and
renders your nice work on the make-up futile, A paper cylinder,
corresponding in calibre to the greatest girth of the birdskin, binds
the wings well, and makes a good specimen. Remarking that there
are some detestable practices, such as hanging up a bird by a string
through the nose (methods only to be mentioned to be condemned),
I will tell you the easiest and best way by which the most elegant
and tasteful results are secured. The skins are simply laid away in
cotton, just as they come from your hands. • Take a considerable
wad of cotton, make a bed of it, lay the specimen in, and tuck
it up nicely around the edges. I generally take a thin sheet of
cotton wadding, the sizing of which confers some textile consist-
ency, and wrap the bird completely but lightly in it. By loosening
or tightening a trifle here or there, laying down a pillow or other
special slight pressure, the most delicate contour- lines may be
preserved with fidelity. Unnecessary pother is sometimes made
about drying skins ; the fact being that under ordinary circumstances
they could not be kept from drying perfectly ; and they dry in

^ Exceptions. Woodpeckers, ducks, and some other birds treated of beyond, are
best set with the bead flat on one side, the bill pointing obliquely to the right or
left ; owls, with the bill pointing straight up in the air as the bird lies on its back.

^ If the mandibles gape, run a thread through the nostrils and tie it tightly under
the bill. Or, since this injures the nostrils (and we frequently want to examine their
structure), stick a pin in under the bill close to the gonys, driving it obliquely into
the palate. Sometimes the skin of the throat looks sunken betwixt the sides of the
jaw. A shred of cotton introduced with forceps through the mouth will obviate
this.

^ Don’t cock up the head, trying to impart a knowing air — it cannot be done, and
only makes the poor bird look ridiculous. Don’t lay the skin on one side, with the
legs in perching position, and don’t spread the wings — the bird will never perch nor
fly again, and the suggestion is not in keeping. The only permissible departure from
the rule of severe simplicity is when some special ornament, as a flne crest, may be
naturally displayed, or some hidden markings be brought out, or a shape of tail or
wing to be perpetuated; but in all such cases the “spread-eagle” style should be
sparingly indulged. It is, however, frequently desirable to give some special set to
hide a defect, as loss of plumage, etc. ; this may often be accomplished very cun-
ningly, with excellent result. No rules for this can be laid down, since the details
vary in every case ; but in general the weak spot may be hidden l3y contracting the
skin of the place, and then setting the bird in an attitude that naturally corresponds,
thus making a virtue of necessity.

SEC. VII

HO IV TO MAKE A BIRD SKIN

51

exactly the shape they are set, if not accidentally pressed upon.
At sea, however, or during unusually protracted wet weather, they
of course dry slowly, and may require some attention to prevent
mildew or souring, especially in the cases of very large, thick-
skinned, or greasy specimens. Thorough poisoning, and drying by
a fire, or placing in the sun, will always answer. Yery close packing
retards drying. When travelling, or operating under other circum-
stances requiring economy of space, you must not expect to turn
out your collection in elegant order. Perfection of contour-lines
can only he secured by putting each specimen away by itself;
undue pressure is always liable to produce unhappily outrS configura-
tion of a skin. Trays in a packing box are of great service in
limiting possibilities of pressure ; they should be shallow ; one four
inches deep will take a well -stuffed hen -hawk, for example, or
accommodate from three to six sparrows atop of one another. It is
well to sort out your specimens somewhat according to size, to keep
heavy ones off little ones ; though the chinks around the former
may usually be economised with advantage by packing in the less
valuable or the less neatly prepared of the latter. When limited
to a travelling chest, I generally pass in the skins as fast as made,
packing them solid in one sense, yet finding a nice resting-place
for each. If each rests in its own cotton cofiin, it is astonishing
how close they may be laid without harm, and how many will go
in a given space ; a tray 30 x 18 x 4 inches will easily hold three
hundred and fifty birds six inches long. As a tray fills up, the
drier ones first put in may be submitted to more pressure. A skin
originally dried in good shape may subsequently be pressed perfectly
flat without material injury ; the only thing to avoid being distor-
tion. The whole knack of packing birds corresponds to that of
filling a trunk solidly full of clothes, as may easily be done without
damage to an immaculate shirt-front. Finally, I would say, never
put away a bird unlabelled, not even for an hour ; you may forget
it or die. Never tie a label to a bird’s bill, wing, or tail ; tie it
securely to legs where they cross, and it will be just half as
liable to become detached as if tied to one leg only. Never paste
a label, or even a number, on a bird’s plumage. Never put in glass
eyes before mounting. Never paint or varnish a bird’s bill or feet.
Never replace missing plumage of one bird with the feathers of
another — no, not even if the birds came out of the same nest.^

^ [In presenting anew, and to an English public, the foregoing directions for mani-
pulation, the author may be pardoned if he alludes to the test of time in their favour.
Some of his earliest specimens, made in 1857, are extant, and in good order still.
Many of the large cabinets, both in Europe and the United States, include some of
his preparations, received in exchange through the Smithsonian Institution, or through
private channels. They will be found, as a rule, compact yet shapely, with a smooth
finish, and very durable. He may add, lest this paragraph should be misunderstood.

52

FIELD ORNITHOLOGY

PART I

(5) Special Processes; Complications and Accidents

The Foregoing’ Method of procedure is a routine practice
applicable to the “general run” of birds. But there are several
cases requiring a modification of this process ; while several circum-
stances may tend to embarrass operations. The principal special
conditions may therefore be separately treated to advantage.

Size. — Other things being equal, a large bird is more difficult
to prepare than a small one. In one case, you only need a certain
delicacy of touch, easily acquired and soon becoming mechanical ;
in the other, demand on your strength may be made, till your
muscles ache. It takes longer, too ; ^ I could put away a dozen
sparrows in the time I should spend over an eagle ; and I would
rather undertake a hundred humming-birds than one ostrich. For
large birds, say anything from a hen-hawk upward, various special
manipulations I have directed may be forgone, while however you
observe their general drift and intent. You may open the bird as
directed, or, turning it tail to you, cut with a knife. ^ Forceps are
rarely required ; there is not much that is too small to be taken in
hand. As soon as the tail is divided, hang up the bird by the

that he has seldom purchased a birdskin, never sold one in his life, and for some
years has owned none. Excepting a few given to friends, his ornithological specimens,
as well as those in other departments of natural history, have always been presented
to the United States Government, and deposited in the national collection at
Washington. 9th September 1889.]

^ The reader may be curious to know something of the statistics on this score —
how long it ought to take him to prepare an ordinary skin. He can scarcely imagine,
from his first tedious operations, how expert he may become, not only in beauty of
result, but in rapidity of execution. I have seen taxidermists make good small skins
at the rate of ten an hour ; but this is extraordinary. The quickest work I ever did
myself was eight an hour, or an average of seven and a half minutes apiece, and fairly
good skins. But 1 picked my birds, all small ones, well shot, labelled, measured,
and plugged beforehand, so that the rate of work was exceptional, besides including
only the actual manipulations from first cut to laying away. No one averages eight
birds an hour, even excluding the necessary preliminaries of cleansing, plugging, etc.
Four birds an hour, everything included, is good work. A very eminent ornithologist
of America, and an expert taxidermist, once laid a whimsical wager that he would
skin and stuff a bird before a certain friend of his could pick all the feathers off a
specimen of the same kind. I forget the time, but he won, and his friend ate crow,
literally, that night.

2 Certain among larger birds are often opened elsewhere than along the belly,
with what advantage I cannot say from my own experience. Various water-birds,
such as loons, grebes, auks, gulls, and ducks (in fact any swimming-bird with dense
under plumage), may be opened along the side by a cut under the wings from the
shoulder over the hip to the rump ; the cut is completely hidden by the make-up,
and the plumage is never ruffled. But I see no necessity for this ; for, as a rule, the
belly-opening can be completely effaced with due care, though a very greasy bird
with white under plumage generally stains where opened, in spite of every precaution.
Such birds as loons, grebes, cormorants, and penguins are often opened by a cut
across the fundament from one leg to the other ; their conformation in fact suggests
and favours this operation. I have often seen water-birds slit down the back ; but I
consider it poor practice.

SEC. VII

HOJV TO MAKE A BIRDSKIN

53

rump, so you will have both hands free. Let it swing clear of the
wall or table, at any height most convenient. The steel hooks of a
dissecting case are not always large enough j use a stout fish-hook
with the barb filed off. Work with your nails, assisted by the
scalpel if necessary. I know of no bird, and I think there is none,
in England at least, the skin of which is so intimately adherent by
fibrous or muscular tissue as to require actual dissecting throughout;
a gannet comes, perhaps, as near this as any ; but in many cases
the knife may be constantly employed with advantage. Use it
with long clean sweeping strokes, hugging the skin rather than the
body. The knee and shoulder commonly require disarticulation,
unless you use bone-nippers or strong shears. To make the four
cuts of the skull may need a very able-bodied instrument, even a
chisel. The wings will give the most trouble, and they require a
special process ; for you cannot readily break up the adhesions of
the secondary quills to the ulna, nor is it desirable that very large
feathers should be deprived of this natural support. Hammer or
nip off the great head of the upper-arm bone, just below the insertion
of the breast-muscles ; clean the rest of that bone and leave it in.
Tie a string around it (what sailors call “ two half-hitches ” gives a
secure hold on the bony cylinder), and tie it to the other humerus,
inside the skin, so that the two bones shall be rather less than their
natural distance apart. After the skin is brought right side out,
attack the wings thus ; Spread the wing under side uppermost, and
secure it on the table by driving a tack or brad through the wrist-
joint ; this fixes the far end, while the weight of the skin steadies
the other. Eaise a whole layer of the under wing-coverts, and make
a cut in the skin thus exposed, from elbow to wrist, in the middle
line between the two fore-arm bones. Eaise the flaps of skin and
all the muscle is laid bare ; it is to be removed. This is best done
by lifting each muscle from its bed separately, slipping the handle of
the scalpel under the individual muscles ; there is little if any bony
attachment except at each end, and this is readily severed. Strew
in arsenic ; a little cotton may be used to fill the bed of muscle
removed from a very large bird ; bring the flaps of skin together,
and smooth down the coverts ; you need not sew up the cut, for
the coverts will hide the opening ; in fact, the operation does not
show at all after the make-up. Stuffing of large birds is not
commonly done with only the four pieces already directed. The
eyeballs, and usually the neck-cylinder, go in as before ; the body
may be filled any way you please, provided you do not put in too
much stufiing nor get any between the shoulders. Large birds had
better have the leg-bones wrapped to nearly natural size. Observe
that the leg -muscles do not form a cylinder, but a cone ; let the
wrapping taper naturally from top to bottom. Attention to this

54

FIELD ORNITHOLOGY

PART I

point is necessary for all large or medium-sized birds with naturally
prominent legs. The stout finely feathered legs of a hawk, for
example, ought to be well displayed ; with these birds, and also
with rails, etc,, moreover, imitate the bulge of the thigh with a
special wad laid inside the skin. Large birds commonly require
also a special wad introduced by the mouth, to make the swell of
the throat ; this wad should be rather fluffy than firm. As a rule,
do not fill out large birds to their natural dimensions ; they take
up too much room. Let the head, neck, and legs be accurately
prepared, but leave the main cavity one-third if not one-half empty;
no more stuffing is required than will fairly smooth out creases in
the skin. Keduce bulk rather by flattening out than by general
compression. Use tow instead of cotton ; and if at all short of tow,
economise with paper, hay, etc., at least for the deeper portions of
the main stuffing. Large birds may be set in a great quantity
of tow ; wrapped in paper, much like any other parcel ; or simply
left to dry on the table, the wings being only supported by
cushioning or other suitable means.

Shape. — Some special configurations have been noticed in the
last paragraph, prematurely perhaps, but leading directly up to
further considerations respecting shape of certain birds as a modify-
ing element in the process of preparation. As for skinning, there
is one extremely important matter. Most ducks, many wood-
peckers, flamingoes, and some others, cannot be skinned in the
usual way, because the head is too large for the calibre of the neck
and cannot be drawn through. In such cases, skin as usual to
the base of the skull, cut off the head there (inside the skin of
course), and operate upon it, after turning the skin right side out,
as follows : Part the feathers carefully in a straight line down the
back of the skull, make a cut through the skin, just long enough to
permit the head to pass, draw out the skull through this opening,
and dress it as already directed. Eeturn it, draw the edges of the
cut nicely together, and sew up the opening with a great many fine
stitches. Simple as it may appear, this process is often embar-
rassing, for the cut has an unhappy tendency to wander about the
neck, enlarging itself even under the most careful manipulation;
while the feathers of the parts are usually so short that it is diffi-
cult to efface all traces of the operation. I consider it very dis-
agreeable ; but for ducks I know of no alternative. I have, however,
found out a way to avoid it with woodpeckers, excepting the very
largest ; it is this : Before skinning, part the eyelids, and plunge the
scalpel right into the eyeball ; seize the cut edge of the ball with
the forceps, and pull the eye right out. It may be dexterously done
without spilling the eye-water on the plumage ; but, for fear of this,
previously put a little gypsum on the spot. Throw arsenic into the

SEC. VII

HO TV TO MAKE A BIRDS KIN

55

socket, and then fill it with cotton poked in between the lids. The
eyes are thus disposed of. Then, in skinning, when you come to
the head, dissever it from the neck and work the skull as far out as
you can j it may be sufficiently exposed, in all cases, for you to
gouge out the base of the skull with the scissors, and get at the
brain to remove it. Apply an extra large dose of arsenic, and you
will never hear from what jaw -muscle has been left in. In all
these cases, as already remarked, the head is preferably set lying
on one side, with the bill pointing obliquely to the right or left.
Certain birds require a special mode of setting ; these are, birds with
very long legs or neck, or both, as swans, geese, pelicans, cormo-
rants, snakebirds, loons, and especially cranes, herons, ibises, and
flamingoes. Long legs should be doubled completely on themselves
by bending at the heel-joint, and either tucked under the wings or
laid on the under surface ; the chief point is to see that the toes lie
flat, so that the claws do not stick up, to catch in things or get
broken off. A long neck should be carefully folded j not at a sharp
angle with a crease in the skin, but with a short curve, and brought
round either to the side of the bird or on its breast, as may seem
most convenient. The object is to make a bale of the skin as
nearly as may be, and when it is properly effected it is surprising
what little space a crane, for instance, occupies. But it is rarely, if
ever, admissible to bend a tail back on the body, however incon-
veniently long it may be. Special dilations of skin, like the pouch of
a pelican, or the air-sacs of a prairie-hen, may be moderately displayed.

Thin Skin. — Loose Plumage. — It is astonishing how much
resistance is offered by the thin skin of the smallest bird. Though
no thicker than tissue paper, it is not very liable to tear if deftly
handled ; yet a rent once started often enlarges to an embarrassing
extent if the skin be stretched in the least. Accidental rents and
enlargements of shot-holes should be neatly sewn up, if occurring in
an exposed place ; but in most cases the plumage may be set to hide
the openings. The trogons are said to have remarkably thin and
delicate skin ; I have never handled one in the flesh. Among
British birds, the species of Caprimulgidce have about the tenderest
skins. The obvious indication in all such cases is simply a little
extra delicacy of manipulation. In skinning most birds, you should
not lose more than a feather or two, excepting those loosened by
the shot. Pigeons are peculiar for the very loose insertion of their
plumage ; you will have to be particularly careful with them, and
in spite of all your precautions a good many feathers will probably
drop. As stripping down the secondary quills from the fore-arm,
in the manner already indicated, will almost invariably set these
feathers free from the skin, I recommend you not to attempt it, but
to dress the wings as prescribed for large birds.

56

FIELD ORNITHOLOGY

PART I

Fatness. — Fat is a substance abhorred of all dissectors ; always
in the way, embarrassing operations and obscuring observations;
while it is seldom worth examination after its structure has once
been ascertained. It is particularly obnoxious to the taxidermist,
since it is liable to soil the plumage during skinning, and also to
soak into the feathers afterwards ; and greasy birdskins are never
pleasing objects. A few birds never seem to have any fat; some,
like petrels, are| always oily ; at times, especially in the indolent
autumn season, when birds have little to do but feed, the great
majority acquire an emlonpoint doubtless to their own satisfaction,
but to the taxidermist’s discomfort. In all such cases gypsum
should be lavishly employed. Strew plaster plentifully from the
first cut all through the operation ; dip your fingers in it frequently,
as well as your instruments. This invaluable absorbent will deal
with most of the running fat. When the skin is completely
reversed, remove as much of the solid fat as possible ; it is generally
found occupying the areolar tissue of particular definite tracts, and
most of it may usually be peeled or flaked off in considerable masses.
Since the soft and oozy state of most birds’ fat at ordinary temper-
atures may be much improved by cold, it will be well to leave your
birds on ice for a while before skinning, if you have the means and
time to do so ; the fat will become quite firm. There is a device
for preventing or at any rate lessening the soiling of the plumage so
apt to occur along the line of incision ; it is invaluable in cases of
white plumage. Take a strip of cloth of greater width than the
length of the feathers, long enough to go up one side of the cut and
down the other. Sew this closely to the skin all around the cut,
and it will form an apron to guard the plumage. You will too
frequently find that a bird, prepared without soiling and laid away
apparently safe, afterwards grows greasy ; if the plumage is white,
it soon becomes worse than ever by showing dust that the grease
catches. Perhaps the majority of such birds in our museums show
the dirty streak along the belly. The reason is, that the grease has
oozed out along the cut, or wherever else the skin has been broken,
and infiltrated the plumage, being drawn up apparently by capillary
attraction, just as a lamp wick sucks up oil. Sometimes, without
obviously soiling the plumage, the grease will run along the thread
that ties the label, and make a uniformly transparent piece of oil-
paper. I have no remedy to offer for this gradual infiltration of
the plumage. It will not wash out, even with soap and water.
Possibly careful and persistent treatment with ether might be
effective, but I am not prepared to say it would be. Eemoval of
all fat that can be got off during skinning, with a liberal use of
plaster, will in a measure prevent a difficulty that remains incurable.

Bloodstains, ete. — In the nature of the case, this complication

SEC. VII

ffOlV TO MAKE A BIRD SKIN

57

is of continual occurrence ; fortunately it is easier dealt with than
greasiness. Much may be done in the field to prevent bloodying of
the plumage, as already said. A little blood does not show much
on a dark plumage ; but it is of course conspicuous on light or
white feathers. Dried blood may often be scraped off, in imitation
of the natural process by which a bird cleanses its plumage with the
bill ; or be pulverised by gently twiddling the feathers between the
fingers, and then blown off. But feathers may by due care be
washed almost as readily as clothing ; and we must ordinarily resort
to this to remove all traces of blood, especially from white surfaces.
If properly dried they do not show the operation. With a soft rag
or pledget of cotton dipped in warm water bathe the place assiduously,
pressing down pretty hard, only taking care to stroke the feathers
the right way, so as not to crumple them, until the red colour dis-
appears ; then you have simply a wet place to deal with. Press
gypsum on the spot ; it will cake ; flake it off and apply more, till
it will no longer stick. Then raise the feathers on a knife -blade
and sprinkle gypsum in among them ; pat it down and shake it up,
till the moisture is entirely absorbed. Two other fluids of the body
will give occasional annoyance, — the juices of the alimentary canal
and the eye-water. Escape of the former by mouth, nostrils, or
vent is preventable by plugging these orifices, and its occurrence is
inexcusable. But shot often lacerates the gullet, crop, and bowels,
and though nothing may flow at the time, subsequent jolting or
pressure in the game-bag causes the escape of fluids : a seemingly
safe specimen may be unwrapped to show the whole belly-plumage
a sodden brown mass. Such accidents should be treated precisely
like bloodstains ; but it is to be remarked that these stains are
not seldom indelible, traces usually persisting, in white plumage at
least, in spite of our best endeavours. Eye-water, insignificant as
it may appear, is often a great annoyance. This liquor is slightly
glairy, or rather glassy, and puts a sort of sizing on the plumage
difficult to efface ; the more so since the soiling necessarily occurs
in a conspicuous place, where the plumage is scanty and delicate.
It frequently happens that a lacerated eyeball, by the elasticity of
the coats, or adhesion of the lids, retains its fluid till this is pressed
out in manipulating the parts ; and, recollecting how the head lies
buried in plumage at that stage of the process, it will be seen that
not only the head, but much of the neck and even the breast, may
become wetted. If the parts are extensively soaked, the specimen
is almost irreparably damaged. Plaster will absorb the moisture,
but much of the sizing may be retained on the plumage ; therefore,
though the place seems simply wet, it should be thoroughly washed
with water before the gypsum is applied. I always endeavour to
prevent the accident ; if I notice a lacerated eyeball, I extract it

58

FIELD ORNITHOLOGY

PART I

before skinning, in the manner described for woodpeckers. Miscel-
laneous stains, from the juices of plants, etc., may be received ; all
such are treated on general principles. Blood on the beak and feet
of rapacious birds, mud on the bill and legs of waders, etc. etc., may
be washed off without the slightest difficulty. A land bird that has
fallen in the water should be recovered as soon as possible, picked
up by the bill, and shaken ; most of the water will run off, unless the
plumage is completely soaked. It should be allowed to dry just as
it is, without touching the plumage, before being wrapped and
bagged. If a bird fall in soft mud, the dirt should be scraped or
snapped off as far as this can be done without plastering the feathers
down, and the rest allowed to dry ; it may afterward be rubbed
fine and dusted off, when no harm will ensue, except to white
feathers, which may require washing.

Mutilation. — You will often be troubled, early in your practice,
with broken legs and wings, and various lacerations; but the injury
must be very severe (such as the carrying away of a limb, or blow-
ing off the whole top of a head) that cannot be in great measure
remedied by care and skill. Suppose a little bird, shot through the
neck or small of the back, comes apart while being skinned ; you
have only to remove the hinder portion, be that much or little, and
go on with the rest as if it were the whole. If the leg-bone of a
small bird be broken near the heel, let it come away altogether ;
it will make little if any difference. In case of the same accident
to a large bird that ought to have the legs wrapped, whittle out a
peg and stick it in the hollow stump of the bone ; if there is no
stump left, file a piece of stout wire to a point and stick it into
the heel joint. If the fore-arm bone that you usually leave in a
small bird is broken, remove it and leave the other in ; if both are
broken, do not clean the wings so thoroughly that they become
detached ; an extra pinch of arsenic will condone the omission. In
a large bird, if both bones of the fore-arm are broken, splint them
with a bit of wood laid in between, so that one end hitches at the
elbow, the other at the wrist. A humerus may be replaced like a
leg bone, but this is rarely required. If the skull be smashed, save
the pieces, and leave them if you can ; if not, imitate the arch of
the head with a firm cotton -ball. A broken tarsus is readily
splinted with a pin thrust up through the sole of the foot ; if too
large for this, use a pointed piece of wire. There is no mending a
bill- when part of it is shot away ; for I think the replacing of part
by putty, stucco, etc., inadmissible ; but if it be only fractured, the
pieces may usually be retained in place by winding with thread, or
with a touch of glue or mucilage. I have already hinted how art-
fully various weak places in a skin, due to mutilation or loss of
plumage, may be hidden.

SEC. VII

HOW TO MAKE A BIRD SKIN

59

Decomposition. — It might seem unnecessary to sjjeak of what
may be smelt so readily as animal putrescence; but there are
some useful points to be learned in this connection, besides the im-
portant sanitary precautions that are to be deduced. Immediately
after death the various fluids of the body begin to settle (so to
speak), and shortly afterward the muscular system becomes fixed
in what is technically called rigor mortis. This stiffening usually
occurs as the animal heat dies away ; but its onset, and especially
its duration, is very variable, according to circumstances, such as
cause of death ; although in most cases of sudden violent death of
an animal in previous good health, it seems to depend chiefly upon
temperature, being transient and imperfect, or altogether wanting,
in hot weather. As it passes off, the whole system relaxes, and the
body soon becomes as limp as at the moment of death. This is the
period immediately preceding decomposition ; in fact, it may be
considered as the stage of incipient putridity ; it is very brief in
warm weather, and it should be seized as the last opportunity of
preparing a bird without inconvenience and even danger. If not
skinned at once, putrescence becomes established ; it is indicated by
the efiluvium; by the distension of the abdomen with gaseous products
of decomposition ; by the loosening of the cuticle, and consequently
of the feathers ; and by other signs. If you part the feathers of a
bad-smelling bird’s belly to find the skin swollen and livid or
greenish, while the feathers come off at a touch, the bird is too far
gone to be recovered without trouble and risk that no ordinary
specimen warrants. It is a singular fact that this early putrescence
is more poisonous than utter rottenness ; as physicians are aware, a
fost-mortem examination at this stage, or even before it, involves
more risk than their ordinary dissecting-room experience. It seems
that both natural and pathological poisons lose their early virulence
by resolution into other products of decay. The obvious deduction
from this is to skin your birds soon enough. Some say they are
best skinned perfectly fresh, but I see no reason for this ; when I
have time to choose, I take the period of rigidity as being prefer-
able on the whole ; for the fluids have then settled, and the
limbs are readily relaxed by manipulation. If you have a large bag
to dispose of, and are pressed for time, set them in the coolest place
you can find, preferably on ice ; a slight lowering of temperature
may make a decided difference. Disembowelling, which may be
accomplished in a moment, will materially retard decomposition.
Injections of creosote or dilute carbolic acid will arrest decay for a
time, or for an indefinitely long period if a large quantity of these
antiseptics be employed. When it becomes desirable (it can never
be necessary) to skin a putrescent bird, great care must be exercised
not only to accomplish the operation, but to avoid danger. I must

6o

FIELD ORNITHOLOGY

PART I

not, however, lead you to exaggerate the risk, and will add that
I think it often overrated. I have probably skinned birds as
“ gamey ” as any one has, and repeatedly, without being conscious
of any ill effects. I am sure that no poison, ordinarily generated
by decomposition of a body healthy at death, can compare in viru-
lence with that commonly resulting after death by many diseases.
I also believe that the gaseous products, however offensive to the
smell, are innocuous as a rule. The danger practically narrows
down to the absorption of fluids through an abraded surface ; the
poison is rarely taken in by natural pores of healthy skin, if it
remain in contact but a short time. Cuts and scratches may be
closed with a film of collodion, or covered with isinglass or court-
plaster, or protected by rubber cots on the fingers. The hands
should, of course, be washed with particular care immediately after
the operation, and the nails scrupulously dressed. Having never
been poisoned, I cannot give the symptoms from personal experience ;
but I will quote from Mr. Maynard : —

“ In a few days numerous pimples, which are exceedingly pain-
ful, appear upon the skin of the face and other parts of the person,
and, upon those parts where there is chafing or rubbing, become
large and deep sores. There is a general languor, and, if badly
poisoned, complete prostration results ; the slightest scratch becomes
a festering sore. Once poisoned in this manner (and I speak from
experience), one is never afterward able to skin any animal that
has become in the least putrid, without experiencing some of the
symptoms above described. Even birds that you handled before
with impunity you cannot now skin without great care. The best
remedy in this case is, as the Hibernian would say, not to get
poisoned . . . bathe the parts frequently in cold water; and, if
chafed, sprinkle the parts after bathing with wheat flour. These
remedies, if persisted in, will effect a cure, if not too bad ; then,
medical advice should be procured without delay.”

My advice would be, to avoid all mechanical irritation of the
inflamed parts ; touch the parts that have ulcerated with a stick of
lunar caustic ; take a dose of salts ; use syrup of the iodide of iron,
or tincture of the chloride of iron, say thirty drops of either, in a
wineglass of water, thrice daily ; rest at first, exercise gradually as
you can bear it; and skin no birds till you have completely
recovered.

How to mount Birds. — As some may not improbably procure
this volume with a reasonable expectation of being taught to mount
birds, I append the required instructions, although I only profess to
treat of the preparation of skins for the cabinet. As a rule, the
purposes of science are best subserved by not mounting specimens ;
for display, the only end attained, is not required. I would

SEC. VII

T^OJV TO MAKE A BIRD SKIN

6i

strongly advise you not to mount your rarer or otherwise par-
ticularly valuable specimens j select for this purpose nice, pretty
birds of no special scientific value. The principal objections to
mounted birds are, that they take up too much room, require special
arrangements for keeping and transportation, and cannot be handled
for study with impunity. Some might suppose that a mounted
bird would give a better idea of its figure and general aspect than a
skin ; but this is only true to a limited extent. Faultless mounting
is an art really difiicult, acquired by few ; the average work done
in this line shows something of caricature, ludicrous or repulsive, as
the case may be. To copy nature faithfully by taxidermy requires
not only long and close study, but an artistic sense ; and this last
is a rare gift. Unless you have at least the germs of the faculty in
your composition, your taxidermal success will be incommensurate
with the time and trouble you bestow. My own taxidermal art is
of a low order, decidedly not above average; although I have
mounted a great many birds that would compare very favourably
with ordinary museum work, few of them have entirely answered
my ideas. A live bird is to me such a beautiful object that the
slightest taxidermal flaw in the effort to represent it is painfully
offensive ; perhaps this makes me place the standard of excellence
too high for practical purposes. I like a good honest birdskin that
does not pretend to be anything else ; it is far preferable to the
ordinary taxidermal abortions of the show-cases. But if, after the
warnings that I mean to . convey in this paragraph, you still wish
to try your hand in the higher department of taxidermy, I will
explain the whole process as far as manipulation goes ; the art you
must discover in yourself.

The operation of skinning is precisely the same as that already
given in detail ; then, instead of stuffing the skin as directed above,
to lie on its back in a drawer, you have to stuff it so that it will
stand up on its feet and look as much like a live bird as possible.
To this end a few additional implements and materials are required.
These are : {a) annealed wire of various numbers ; it may be iron,
copper, or brass, but must be perfectly annealed, so as to retain no
elasticity or spring ; (5) several files of different sizes ; (c) some
slender straight brad-awls ; (cZ) cutting pliers ; (e) setting needles,
merely sewing or darning needles stuck in a light wooden handle,
for dressing individual feathers ; (/) plenty of pins (the long, slender
insect pins used by entomologists are the best) and sewing thread ;
(y) an assortment of glass eyes. (The fixtures and decorations are
noticed, beyond, as occasion for their use arises.)

There are two principal methods of mounting, which may be
respectively styled soft stuffing and hard stuffing. In the former, a
wire framework, consisting of a single anterior piece passing in the

62

FIELD ORNITHOLOGY

PART I

middle line of the body up through the neck and out at the top of
the head, is immovably joined behind with two pieces, one passing
through each leg ; around this naked forked frame soft stuffing is
introduced, bit by bit, till the proper contour of the skin is secured.
I have seen very pretty work of this kind, particularly on small
birds ; but I consider it much more difficult to secure satisfactory
results in this way than by hard stuffing, and I shall therefore con-
fine attention to the latter. This method is applicable to all birds,
is readily practised, facilitates setting of the wings, arranging of the
plumage, and giving of any desired attitude. In hard stuffing, you
make a firm ball of tow rolled upon a wire of the size and shape of
the bird’s body and neck together ; you introduce this whole, after-
wards running in the leg wires and clinching them immovably in
the mass of tow.

Having your empty skin in good shape, as already described,
cut three pieces of wire of the right ^ size ; one piece somewhat
longer than the whole bird, the other pieces two or three times as
long as the whole leg of the bird. File one end of each piece to a
fine sharp point; try to secure a three -edged cutting point like
that of a surgical needle, rather than the smooth punching point
of a sewing-needle, as the former perforates more readily. Have
these wires perfectly straight.^ Bend a small portion of the unfiled
end of the longer wire irregularly upon itself, as a convenient
nucleus for the ball of tow.^ Take fine clean tow, in loose dossils,
and wrap it round and round the wire nucleus, till you make a firm
ball, of the size and shape of the bird’s body and neck. Study the
contour of the skinned body : notice the swelling breast-muscles,
the arch of the lower back, the hollow between the forks of the
merrythought into which the neck, when naturally curved, sinks.
Everything depends upon correct shaping of the artificial body ; if
it be misshapen, no art can properly adjust the skin over it. Firm-
ness of the tow ball and accurate contour may both be secured by
wrapping the mass with sewing thread, loosening here, tightening
there, till the shape is satisfactory. Be particular to secure a smooth
surface ; the skin in drying will shrink close to the stuffing, dis-
closing its irregularities, if there be any, by the maladjustment of
the plumage that will ensue. Observe especially that the neck,
though the direct continuation of the backbone, dips at its lower

1 The right size is the smallest that will support the whole weight of the stuffing
and skin without bending, when a piece is introduced into each leg. If using too
thick wire, you may have trouble in thrusting it through the legs, or may burst the
tarsal envelope.

2 If accidentally kinky, the finer sizes of wire may be readily straightened by
drawing strongly upon them so as to stretch them a little. Heavier wire must be
hammered out straight.

2 Cotton will not do at all ; it is too soft and elastic, and moreover will not allow
of the leg wires being thrust into it and there clinched.

SEC. VII

BOW TO MAKE A BIRD SKIN

63

end into the hollow of the merrythought, and so virtually begins
there instead of directly between the shoulders. The three mis-
takes most likely to be made by a beginner are, getting the body
altogether too large, not firm enough, and irregular. When properly
made, it will closely • resemble the bird’s body and neck, with an
inch or several inches of sharp -pointed wire protruding from
the anterior extremity of the neck of tow. You have now to
introduce the whole affair into the skin. With the birdskin on its
back, the tail pointing to your right elbow, and the abdominal
opening as wide as possible, hold the tow body in position relative
to the skin ; enter the wire, pass it up through the neck, bring the
sharp point exactly against the middle of the skull, pierce skull and
skin, causing the wire to protrude some distance from the middle
of the crown. Then by gentle means insinuate the body, partly
pushing it in, partly drawing the skin over it, till it rests in its
proper position. This is just like drawing on a tight kid glove, and
no more difficult. See that the body is completely encased ; you
must be able to close the abdominal aperture entirely. You have
next to wire the legs. Enter the sharp point of one of the leg-
wires already prepared, exactly at the centre of the sole of the foot,
thrusting it up inside the tarsal envelope the whole length of the
shank, thence across the heel-joint ^ and up along the next bone
of the leg, still inside the skin. The point of the wire will then be
seen within the skin, and may be seized and drawn a little farther
through, and you will have passed a wire entirely out of sight all
the way along the leg. The end of the wire is next to be fixed
immovably in the tow ball. Thrust it in at the point where the
knee, in life, rests against the side of the body.^ Bring the point
to view, bend it over and reinsert it till it sticks fast. There are no
special directions to be given here ; fasten the wire in any way that
effectually prevents wabbling. You may find it convenient to
wire both legs before fastening either, and then clinch them by
twisting the two ends together. But remember that the leg-wires
may be fixed respecting each other, yet permit a see-saw motion of
the body upon them. This must not be ; the body and legs must
be fixed upon a jointless frame. Having secured the legs, close the
abdominal opening nicely, either by sewing or pinning ; you may
stick pins in anywhere, as freely as in a pin-cushion ; the feathers

^ There is occasionally difficulty in getting the wire across this joint, from the
point sticking into the enlarged end of the shin-hone. In such case, take stout
pliers and pinch the joint till the hone is smashed to fragments. The wire will then
pass and the comminution will not show. If there is any trouble in passing the
wire through the tarsus, hore a hole for it with a hrad-awl.

^ This point is farther forward and more helly-ward than you might suppose.
Observe the skinned body again, and see where the lower end of the thigh lies. If
you insert the wire too far back, you cannot by any possibility balance the bird
naturally on its perch ; it will look in imminent danger of toppling over.

64

FIELD ORNITHOLOGY

PART I

hide their heads. Stick a pin through the pope’s-nose to fix the
tail in place.

All this while the bird has been lying on its back, the neck
stretched straight in continuation of the body, wired stiffly, the legs
straddling wide apart, straight and stiff*, the wings lying loosely,
half-spread. Now bring the legs together, parallel with each other,
and make the sharp bend at the heel-joint that will bring the feet
naturally under the belly (over it, as the bird lies on its back).
Pick up the bird by the wires that project from the soles and set it
on its stand, by running the wires through holes bored the proper
distance apart, and then securing the ends by twisting. The
temporary stand that you use for this purpose should have a heavy
or otherwise firm support, so as not easily to overturn during the
subsequent manipulations. At this stage the bird is a sorry-looking
object \ but if you have stuffed correctly and wired securely, it will
soon improve. Begin by making it stand properly. The common
fault here is placing the tarsi too nearly perpendicular. Perching
birds, constituting the majority, habitually stand with the tarsi more
nearly horizontal than perpendicular, and generally keep the tarsi
parallel with each other. Wading and most walking birds stand
with the legs more nearly upright and straight. Many swimming
birds straddle a little ; others rarely if ever. See that the toes
clasp the perch naturally, or are properly spread on the flat surface.
Cause the flank feathers to be correctly adjusted over the tibise (and
here I will remark that with most birds little, if any, of the tibise
shows in life), the heel-joint barely, if at all, projecting from the
general plumage. It is a common fault of stuffing not to draw the
legs closely enough to the body. Above all, look out for the centre
of gravity ; though you have really fastened the bird to its perch,
you must not let it look as if it would fall off if the wires slipped ;
it must appear to rest there of its own accord. Next, give the head
and neck a preliminary setting, according to the attitude you have
determined upon. This will bring the plumage about the shoulders
in proper position for the setting of the wings, to which you may
at once attend. If the body be correctly fashioned and the skin of
the shoulders duly adjusted over it, the wings will fold into place
without the slightest difficulty. All that I have said before about
setting the wings in a skin applies here as well j but in this case
they will not stay in place, since they fall by their own weight.
They must be pinned up. Holding the wing in place, thrust a pin
steadily through near the wrist-joint, into the tow body. Some-
times another pin is required to support the weight of the primaries ;
it may be stuck into the ffank of the bird, the outer quill feather
resting directly upon it. With large birds a sharp pointed wire
must replace the pin. When properly set, the wing-tips will fall

SEC. VII

JIOfF TO MAKE A BIRD SKIN

65

together or symmetrically opposite each other, the quills and
coverts will he smoothly imbricated, the scapular series of feathers
will lie close, and no bare space will show in front of the shoulder.
Much depends upon the final adjustment of the head. The
commonest mistake is getting it too far away from the body. In
the ordinary attitudes of most birds little neck shows, the head
appearing nestled upon the shoulders. If the neck appears too
long, it is not to be contracted by pushing the head directly down
upon it, but by making an S curve of the neck. No precise
directions can be given for the set of the head, but you may be
assured it is a delicate, difficult matter ; the slightest turn of the
bill one way or another may alter the whole expression of the bird.
You will of course have determined beforehand upon your attitude,
upon what you wish the bird to appear to be doing ; then, let your
meaning be pointed by the bird’s bill.

On the general subject of striking an attitude, and giving
expression to a stuffed bird, little can be said to good purpose. If
you are to become proficient in this art, it will come from your
own study of birds in the field, your own good taste and apprecia-
tion of bird -life. The manual processes are easily described and
practised ; it is easy to grind paint, I suppose, but not so to be an
artist. I shall therefore only follow the above account of the
general processes with some special practical points. After “at-
titudinising ” to your satisfaction, or to the best of your ability, the
plumage is to be carefully dressed. Feathers awry may be set
in place with a light spring forceps, or needles fixed in a handle,
one by one if necessary. When no individual feather seems out of
place, it often occurs that the general plumage has a loose, slovenly
aspect. This is readily corrected by wrapping with fine thread.
Stick a pin into the middle of the back, another into the breast,
and perhaps others elsewhere. Fasten the end of a spool of sewing
cotton to one of the pins, and carry it to another, winding the
thread about among the pins, till the whole surface is covered with
an irregular network. Tighten to reduce an undue prominence,
loosen over a depression ; but let the wrapping as a whole be light,
firm, and even. This procedure, nicely executed, will give a smooth-
ness to the plumage not otherwise attainable, and may be made to
produce the most exquisite curves, particularly about the head,
neck, and breast. The thread should be left on till the bird is
perfectly dry ; it may then be unwound or cut off, and the pins
withdrawn. When a particular patch of skin is out of place, it
may often be pulled into position and pinned there. You need not
be afraid of sticking pins in anywhere : they may be buried in the
plumage and left there, or withdrawn when the skin is dry. In
addition to the main stuffing, a little is often required in particular

66

FIELD ORNITHOLOGY

PART I

places. As for the legs, they should be filled out in all such cases
as I indicated earlier in this section ; small birds require no such
stuffing. It is necessary to fill out the eyes so that the lids rest
naturally ; it may be done as heretofore directed, or by putting in
pledgets of cotton from the outside. A little nice stuffing is gener-
ally required about the upper throat.

To stuff a bird with spread wings requires a special process, in
most cases. The wings are to be wired, exactly as directed for the
legs ; they may then be placed in any shape. But with most small
birds, and those with short wings, simple pinning in the half-spread
position indicating fluttering will suffice ; it is readily accomplished
with a long, slender insect pin. I have already spoken of fixing
the tail by pinning or wiring the pope’s-nose to the tow body ;
it may be thus fixed at any desired elevation or depression. There
are two ways of spreading the tail. One is to run a pointed wire
through the quills, near their base, where the wire will be hidden
by the coverts ; each feather may be set at any required distance
from the next by sliding it along this wire. This method is appli-
cable to large birds ; for small ones the tail may be fixed with the
desired spread by enclosing it near its base in a split match, or two
slips of cardboard, with the ends tied together. This holds the
feathers until they dry in position, when it is to be taken off.
Crests may be raised, spread, and displayed on similar principles.
A small crest, like that of a cardinal or cherry bird, for instance,
may be held up till it dries in position by sticking in behind it a
pin with a little ball of cotton on its head. It is sometimes neces-
sary to make a bird’s toes grasp a support by tying them down to
it till they dry. The toes of waders that do not lie evenly on the
surface of the stand may be tacked down with small brads. The
bill may be pinned open or shut, as desired, by the method already
given.

Substitution of an artificial eye for the natural one is essential
for the good looks of a specimen. Glass eyes, of all sizes and
colours, may be purchased at a moderate cost. The pupil is always
black ; the iris varies. You will, of course, secure the proper colour
if it is known, but if not, put in a dark brown or black eye. It is
well understood that this means nothing ; it is purely conventional.
Yellow is probably the next most common colour ; then come red,
white, blue, and green, perhaps approximately in this order of
frequency. But do not use these striking colours at haphazard ;
sacrificing truth, perhaps, to looks. Eyes are generally inserted
after the specimen is dry. Remove a portion of the cotton from
the orbit, and moisten the lids till they are perfectly pliable ; fix
the eye in with putty or wet plaster of Paris, making sure that the
lids are naturally adjusted over it. It goes in obliquely, like a

SEC. VII

HOW TO MAKE A BIRD SKIN

67

button through a button-hole. Much art may be displayed in this
little matter, making a bird look this way or that, to carry out the
general expression.

On finishing a specimen, set it away to dry ; the time required
varies, of course, with the weather, the size of the bird, its fatness,
etc. The more slowly it dries, the better ; there is less risk of the
skin shrinking irregularly. You will often find that a specimen set
away with smooth plumage and satisfactory curves dries more or
less out of shape, perhaps with the feathers raised in places. I
know of no remedy ; it may, in a measure, be prevented by scrupu-
lous care in making the body smooth and firm, and in securing
slow, equable drying. When perfectly dry remove the wrapping,
pull out the superfluous pins or wires, nip off the others so short
that the ends are concealed, and insert the eyes. The specimen is
then ready to be transferred to its permanent stand.

Fixtures for the display of the object of course vary intermin-
ably. We will take the simplest case, of a large collection of
mounted birds for public exhibition. In this instance, uniformity
and simplicity are desirable. “ Spread eagle ” styles of mounting,
artificial rocks and flowers, etc., are entirely out of place in a
collection of any scientific pretensions, or designed for popular
instruction. Besides, they take up too much room. Artistic group-
ing of an extensive collection is usually out of the question ; and
when this is unattainable, half-way efforts in that direction should
be abandoned in favour of severe simplicity. Birds look best on
the whole in uniform rows, assorted according to size, as far as a
natural classification allows. They are best set on the plainest
stands, with circular base and a short cylindrical crossbar on a
lightly turned upright. The stands should be painted dead- white,
and be no larger than is necessary for secure support ; a neat stiff
paper label may be attached. A small collection of birds, as an
ornament to a private residence, offers a different case ; here variety
of attitude and appropriate imitation of the birds’ natural surround-
ings are to be secured. A miniature tree, on which a number of
birds may be placed, is readily made. Take stout wire, and by
bending it, and attaching other pieces, get the framework of the
tree of the desired size, shape, and number of perches. Wrap it
closely with tow to a proper calibre, remembering that the two
forks of a stem must be together only about as large as the stem
itself. Gather a basketful of lichens and tree moss ; reduce them
to coarse powder by rubbing with the hands ; besmear the whole
tree with mucilage or thin glue, and sift the lichen powder on it
till the tow is completely hidden. This produces a very natural
effect, which may be heightened by separately affixing larger scraps
of lichen, or little bunches of moss j artificial leaves and flowers

68

FIELD ORNITHOLOGY

PART I

may be added at your taste. The groundwork may be similarly
prepared with a bit of board, made adhesive and bestrewn with the
same substance ; grasses and moss may be added. If a flat surface
is not desired, soak stout pasteboard till it can be moulded in
various irregular elevations and depressions ; lay it over the board
and decorate it in the same way. Kocks may be thus nicely
imitated, with the addition of powdered glass of various colours.
Such a lot of birds is generally enclosed in a cylindrical glass case
with arched top. As it stands on a table to be viewed from
different points, it must be presentable on all sides. A niche in
parlour or study is often fitted with a wall -case, which, when
artistically arranged, has a very pleasing, effect. As such cases may
be of considerable size, there is opportunity for the display of
great taste in grouping. A place is not to be found for a bird, but
a bird for the place, — waders and swimmers below on the ground,
perchers on projecting rests above. The surroundings may be
prepared by the methods just indicated. One point deserves atten-
tion here : since the birds are only viewed from the front, they
may have a “ show-side ” to which everything else may be sacrificed.
Birds are represented flying in such cases more readily than under
other circumstances, supported on a concealed wire inserted in the
back of the case. I have seen some very successful attempts to
represent a bird swimming, the duck being let down part way
through an oval hole in a plate of thick glass, underneath which
were fixed stuffed fishes, shells, and seaweed. It is hardly necessary
to add that in all ornamental collections, labels or other scientific
machinery must be rigorously suppressed.

Transportation of mounted birds offers obvious difficulty. Un-
less very small, they are best secured immovably inside a box by
screwing the foot of the stands to the bottom and sides, so that
they stay in place without touching each other. Or, they may be
carefully packed in cotton, with or without removal of the stands.
Their preservation from accidental injury depends upon the same
care that is bestowed upon ordinary fragile .ornaments of the
parlour. The ravages of insects are to be prevented upon the
principles to be hereafter given in treating of the preservation of
birdskins.

§ 8.— MISCELLANEOUS PAETICULABS

Determination of Sex. — This is an important matter, which
should never be neglected. For although many birds show un-
equivocal sexual distinctions of size, shape, and colour, like those of

SEC. VIII

MISCELLANEOUS PARTICULARS

69

the barnyard cock and hen, for instance, yet the outward character-
istics are more frequently obscure, if not altogether inappreciable,
on examination of the skin alone. Young birds, moreover, are
usually indistinguishable as to sex, although the adults of the same
species may be easily recognised. The rule results, that the sexual
organs should be examined as the only infallible indices. The
essential organs of masculinity are the testicles ; similarly, the ovaries
contain the essence of the female nature. However similar the
accessory sexual structures may be, the testicles and ovaries are
always distinct. The male organs of birds never leave the cavity
of the belly to fill an external bag of skin (scrotum) as they do
among mammals ; they remain within the abdomen, and lie in the
same position as the ovaries of the female. Both these organs are
situated in the belly opposite the “ small of the back,” bound
closely to the spine, resting on the front of the kidneys near their
fore end. The testicles are a pair of subspherical or rather ellip-
soidal bodies, usually of the same size, shape, and colour, and are
commonly of a dull opaque whitish tint. They always lie close
together. A remarkable fact connected with them is, that they are
not always of the same size in the same bird, being subject to
periodical enlargement during the breeding season, and corresponding
atrophy at other seasons. Thus the testicles of a house sparrow,
no bigger than a pin’s head in winter, swell to the size of peas in
April. The ovary (for although this organ is paired originally, only
one is usually functionally developed in birds) will be recognised as a
flattish mass of irregular contour, and usually whitish colour ; when
inactive, it simply appears of finely granular structure which may
require a hand lens to be made out ; when producing eggs, its
appearance is unmistakable. Both testis and ovary may further be
recognised by a thread leading to the end of the lower bowel, — in
one case the sperm-duct, in the other the oviduct ; the latter is
usually much the more conspicuous, as it at times transmits the
perfect egg. There is no difficulty in reaching the site of these
organs. Lay the bird on the left side, its belly toward you : cut
with the scissors through the belly-walls diagonally from anus to
the root of the last rib, or further, snipping across a few of the lower
ribs, if these continue far down, as they do in a loon, for instance.
Press the whole mass of intestines aside collectively, and you at
once see to the small of the back. There you observe the Tddneys,
— large, lobular, dark reddish masses moulded into the concavity
of the sacrum (or back middle bone of the pelvis) ; and on their
surface, toward their fore end, lie testes or ovary, as just described.
The only precaution required is, not to mistake for testicles a pair
of small bodies capping the kidneys. These are the adrenals or
suprarenal capsules, — organs whose function is unknown, but

70

FIELD ORNITHOLOGY

PART I

with which at any rate we have nothing to do in this connection.
They occur in both sexes, and if the testicles are not immediately
seen, or the ovary not at once recognised, they might easily be
mistaken for testicles. Observe that, instead of lying in front, they
cap the kidneys ; that they are usually yellowish instead of opaque
whitish ; and that they have not the firm, smooth, regular sphericity
of the testicles. The testes, however, vary more in shape and
colour than might be expected, being sometimes rather oblong or
linear, and sometimes grayish or livid bluish, or reddish. There is
occasionally but one. The sex determined, use the sign c? or $ to
designate it, as already explained.

Reeog’nition of Age is a matter of ornithological experience
requiring in many or most cases great familiarity with birds for its
even approximate accomplishment. There are, however, some un-
mistakable signs of immaturity, even after a bird has become full-
feathered, that persist for at least one season. These are, in the
first place, a peculiar soft fluffy feel of the plumage ; the feathers
lack a certain smoothness, density, and stiffening which they subse-
quently acquire. Secondly, the bill and feet are softer than those
of the adults ; the corners of the mouth are puffy and flabby, the
edges and point of the bill are dull, and the scales, etc., of the legs
are not sharply cut. Thirdly, the flesh itself is tender and pale
coloured. These are some of the points common to all birds, and
are independent of the special markings that belong to the youth of
particular species. Some birds are actually larger for a while after
leaving the nest than in after years when the frame seems to shrink
somewhat in acquiring the compactness of senility. On the other
hand, the various members, especially the bill and feet, are propor-
tionally smaller at first. Newly growing quills are usually recog-
nised on sight, the barrel being dark coloured and full of liquid,
while the vanes are incomplete. In studying, for example, the
shape of a wing or tail, there is always reason to suspect that the
natural proportions are not yet presented, unless the quill is dry,
colourless, and empty, or only occupied with shrunken white pith.

Examination of the Stomach frequently leads to interesting
observations, and is always worth while. In the first place, we
learn most unquestionably the nature of the bird’s food, which is a
highly important item in its natural history. Secondly, we often
secure valuable specimens in other departments of zoology, particu-
larly entomology. Birds consume incalculable numbers of insects,
the harder kinds of which, such as beetles, are not seldom found
intact in their stomachs j and- a due percentage of these represent
rare and curious species. The gizzards of birds of prey, in particular,
should always be inspected, in search of the small mammals, etc.,
they devour ; and even if the creatures are unfit for preservation.

SEC. VIII

MISCELLANEOUS PARTICULARS

r

we at least learn of their occurrence, perhaps unknown before in a
particular region. Mollusk-feeding and fish-eating birds yield their
share of specimens. The alimentary canal is often the seat of
parasites of various kinds, interesting to the helminthologist ; other
species are to be found under the skin, in the body of muscle, in
the brain, etc. Most birds are also infested with external parasites
of many kinds, so various that almost every leading species has its
own sort of louse, tick, etc. Since these creatures are only at home
with a live host, they will be found crawling on the surface of the
plumage, preparing for departure, as soon as the body cools after
death. There is thus much to learn of a bird aside from what the
prepared specimen teaches, and moreover apart from regular ana-
tomical investigations. Whenever practicable, brief items should
be recorded on the label, as already mentioned.

Restoration of Poor Skins. — If your cabinet be a general
one, comprising specimens from various sources, you will frequently
happen to receive skins so badly prepared as to be unpleasant
objects, besides failing to show their specific characters. There
is, of course, no supplying of missing parts or plumage ; but if
the defect be simply deformity, this may usually be in a measure
remedied. The point is simply to relax the skin, and then proceed
as if it were freshly removed from the bird ; it is what bird-stuffers
constantly do in mounting birds from prepared skins. The relaxa-
tion is effected by moisture alone. Remove the stuffing; fill the
interior with cotton or tow saturated with water, yet not dripping ;
put pads of the same under the wings ; wrap the bill and feet, and
set the specimen in a damp, cool place. Small birds soften very
readily and completely ; the process may be facilitated by persistent
manipulation. This is the usual method, but there is another,
more thorough and more effective ; it is exposure to a vapour-bath.
The appointments of the kitchen stove furnish all the apparatus
required for an extempore steamer; the regular fixture is a tin
vessel much like a wash-boiler, with closed lid, false bottom, and
stopcock at lower edge. On the false bottom is placed a heavy
layer of gypsum, completely saturated with water ; the birds are
laid on a perforated tray above it ; and a gentle heat is maintained
over a stove. The vapour penetrates every part of the skin, and
completely relaxes it, without actually wetting the feathers. The
time required varies greatly of course ; observation is the best
guide. The chief precaution is not to let the thing get too hot.
Professor Baird has remarked that crumpled or bent feathers may
have much of their original elasticity restored by dipping in hot
water. Immersion for a few seconds suffices, when the feathers
will be observed to straighten out. Shaking off superfluous water,
they may be simply left to dry, or they may be dried with plaster.

72

FIELD ORNITHOLOGY

PART I

The method is chiefly aj^plicahle to the large feathers of the wings
and tail. Soiled plumage of dried skins may be treated exactly as
in the case of fresh skins.

Mummification. — As before mentioned, decay may be arrested
by injections of carbolic acid and other antiseptics ; if the tissues be
sufficiently permeated with these substances, the body will keep
indefinitely; it dries and hardens, becoming, in short, a mummy.
Injection should be done by the mouth and vent, be thorough, and
be repeated several times as the fluid dries in. It is an improve-
ment on this to disembowel and fill the belly with saturated
tow or cotton. Due care should be taken not to soil the feathers
in any case, nor should the carbolic solution come in contact with
the hands, for it is a powerful irritant poison. I mention the
process chiefly to condemn it ; I cannot imagine what circumstances
would recommend it, while only an extreme emergency could
justify it. It is further objectionable because it appears to lend a
dingy hue to some plumages, and to dull most of them perceptibly.
Birds prepared — rather unprepared — in this way, may be relaxed
by the method just described, and then skinned ; but the operation
is difficult.

Wet Preparations. — By this term is technically understood an
object immersed in some preservative fluid. It is highly desirable
to obtain more information of birds than their stuffed skins can ever
furnish, and their structure cannot be always examined by dissection
on the spot. In fact, a certain small proportion of the birds of any
extensive collecting may be preferably and very profitably preserved
in this way. Specimens in too poor plumage to be worth skinning
may be thus utilised ; so may the bodies of skinned birds, which,
although necessarily defective, retain all the viscera, and also afford
osteological material. Alcohol is the liquid usually employed, and,
of all the various articles recommended, seems to answer best on
the whole. I have used a very weak solution of chloride of zinc
with excellent results ; it should not be strong enough to show the
slightest turbidity. As glass bottles are liable to break when
travelling, do not fit corners, and offer practical annoyance about
corkage, rectangular metal cans, preferably of copper, with screw-
lid opening, are advisable. They are to be set in small, strong,
wooden boxes, made to leave a little room for the lid -wrench,
muslin bags for doing up separate parcels, parchment for labels, etc.
Unoccupied space in the cans should be filled with tow or a
similar substance, to prevent the specimens from swashing about.
Labelling should be on parchment ; the writing should be perfectly
dry before immersion ; india-ink is the best. Skinned bodies
should be numbered to correspond with the dried skin from
which taken ; otherwise they may not be identifiable. Large birds

SEC. VIII

MISCELLANEOUS PARTICULARS

73

thrown in unskinned should have the belly opened, to let in the
alcohol freely. Birds may be skinned, after being in alcohol, by
simply drying them ; they often make fair specimens. Watery
moisture that may remain after evaporation of the alcohol may
be dried with plaster.

Osteologieal and other Preparations (Figs. 1-3). — While complete
skeletonising of a bird is a special art of some difficulty, and one
that does not fall within the scope of this treatise, I may mention
two bony preparations very readily made, and capable of rendering
ornithology essential service. I refer to the skull, and to the
breast -bone with its principal attachments. These parts of the
skeleton are, as a rule, so highly characteristic that they afford in
most cases invaluable zoological items. To save a skull is of course
to sacrifice a skin, to all intents ; but you often have mutilated or

Figs. 1, 2. — Views of sternum and pectoral arch of the ptarmigan, Lagopus albus, reduced ;
after A. Newton. (I) Lateral view, with the hones upside down; (2) viewed from below, a,
sternum or breast-bone, showing two long slender lateral processes ; h, ends of sternal ribs ; c,
ends of humerus, or upper-arm bone, near the shoulder-joint ; d, scapula, or shoulder-blade ;
e, coracoid ; /, merrythought, or furculum (clavicles).

decayed specimens that are very profitably utilised in tliis way.
The breast-bone (Figs. 1, 2, a) excepting when mutilated, is always
preservable with the skin, and for choice invoices may form its
natural accompaniment. You want to remove along with it the
coracoids (the stout bones connecting the breast -bone with the
shoulders. Figs. 1, 2, e), the merrythought (Figs. 1, 2,/) intervening
between these bones, and the shoulder-blades (Figs. 1, 2, d), all
without detachment from each other, for these bones collectively
constitute the shoulder-girdle, or scapidar arch. Slice off the large
breast muscles close to the bone, and divide their insertions into
the wing-bones (c) ; scrape or cut away the muscles that tie the
shoulder-blades to the chest ; snip off the ribs (Figs. 1, 2, h) close to
the side of the breast-bone ; sever a tough membrane usually found
between the prongs of the wish-bone ; then, by taking hold of the
shoulders (Figs. 1, 2, at c), you can lift out the whole affair, dividing
some slight connections underneath the bone and behind it. The

74

FIELD ORNITHOLOGY

PART I

following points require attention ; the breast-bone often has long
slender processes behind and on the sides (the common fowl and the
ptarmigan are extreme illustrations of this, as shown in the figures),
liable to be cut by mistake for ribs, or to be snapped \ the shoulder-
blades usually taper to a point, easily broken off ; the merrythought
is sometimes very delicate or defective. When travelling, it is
generally not advisable to make perfect preparations of either skull
or sternum ; they are best dried with only superfluous flesh removed,
and besprinkled with arsenic. The skull, if perfectly cleaned, is
particularly liable to lose the anvil-shaped, pronged bones that hinge
the jaw, and the freely movable pair that push on the palate from
behind. Great care should be exercised respecting the identification
of these bones, particularly the sternum, which should invariably
bear the number of the specimen to which it belongs ; the label
should be tied to the coracoid bone. A skull is more likely to be
able to speak for itself, and, besides, is not usually accompanied by
a skin ; nevertheless, any record tending to facilitate its recognition
should be duly entered on the register. There are methods of
making elegant bony preparations. You may secure very good
results by simply boiling the bones, or, what is perhaps better,
macerating them in water till the flesh is completely rotted away,
and then bleaching them in the sun. A little potassa or soda
hastens the process. With breast-bones, if you can stop the process
just when the flesh is completely dissolved, but the tougher liga-
ments remain, you secure a “natural” preparation, as it is called;
if the ligaments go too, the associate parts of a large specimen may
be wired together, those of a small one glued. I think it best,
with skulls, to clean them entirely of ligament as well as muscle ;
for the underneath parts are usually those conveying the most
desirable information, and they should not bo in the slightest
degree obscured. Since in such case the anvil-shaped bones, the
palatal cylinders already mentioned, and sometimes other portions
come apart, the whole are best kept in a suitable box. I prefer to
see a skull with the sheath of the beak removed, though in some
cases, particularly of hard-billed birds, it may profitably be left on.
The completed preparations should be fully labelled by writing on
the bone, in preference to an accompanying or attached paper slip,
which may be lost. Some object to this, as others do to writing on
eggs, that it defaces the specimen ; but I confess I see in dry
bones no beauty but that of utility.

“ In many families of birds, as the ducks {Anatidoi), the trachea
or windpipe of the male affords valuable means of distinguishing
between the different natural groups, or even species, chiefly by the
form of the bony labyrinth, or bulla ossea, situated at or just above
the divarication of the bronchial tubes. A little trouble will enable

SEC. IX

COLLECTION OF NESTS AND EGGS

75

the collector in all cases to preserve this organ perfectly, as repre-
sented in the annexed engraving (Fig. 3). Before proceeding to
skin the specimen a narrow-hladed knife should
be introduced into its mouth, and by taking
hold of the tongue (A) by the fingers or for-
ceps, the muscles (B B) by which it is attached
to the lower jaw should be severed as far as
they can be reached, care being of course taken
not to puncture the windpipe {C C) ; and later
in the operation of skinning, when dividing
the body from the neck or head, not to cut
into or through it. This done, the windpipe
can be easily withdrawn entire and separated
from the neck, and then the sternal apparatus
being removed as before described, its course
must be traced to where, after branching off
in a fork (D), the bronchial tubes (E E) join
the lungs. At these latter points it is to be
cut off. Then rinsing it in cold water, and
leaving it to dry partially, it may, while yet
pliant, be either wrapped round the sternum,
or coiled up and labelled separately” (Professor
Alfred Newton).

§ 9.— COLLECTION OF NESTS
AND EGGS

Opnithology and Oology are twin studies,
or rather one includes the other. A collection
of nests and eggs is indispensable for any
thorough study of birds ; and many persons
find peculiar pleasure in forming one. Some,
however, shrink from robbing birds’ nests as
something particularly cruel — a sentiment
springing, no doubt, from the sympathy and
deference that the tender office of maternity
inspires. But with all proper respect for the
humane emotion, it may be said simply, that
birds’-nesting is not nearly so cruel as bird-
shooting. What I said in a former section,
in endeavouring to guide search for birds,
applies in substance to hunting for their nests ;
the essential difference is, that the latter are of

Fig. 3. — Trachea or wind-
pipe of the male red-breasted
merganser, Mergus serrator,
about J nat. size, viewed from
above (behind) ; after Newton.
A, tongue ; B B, its attach-
ments ; C C, windpipe, dilated
in the middle and swelling
below into a bony box, D ;
E E, bronchial tubes, going
to lungs.

76

FIELD ORNITHOLOGY

PART I

course stationary objects, and consequently more liable to be over-
looked, other things being equal, than birds themselves. Most birds
nest on trees or bushes ; many on the ground and on rocks ; others in
hollows. Some build elegant, elaborate structures, endlessly varied
in details of form and material ; others make no nest whatever.
Egging is chiefly practicable in May and during the summer ; but
some species, particularly birds of prey, begin to lay late in winter
or early in spring, so there is really a long period for search. Par-
ticular nests, of course, like the birds that build them, can only be
found through ornithological knowledge ; but general search is
usually rewarded with a varied assortment. The best clue to a
hidden nest is the action of the parents ; patient watchfulness is
commonly successful in tracing the bird’s home. As the science of
oology has not progressed to the point of determining from the nests
and eggs to what bird they belong, in even a majority of cases, the
utmost care in authentication is indispensable. To be worth anything,
not to be worse than worthless in fact, an egg must be identified be-
yond question ; must be not only unsuspected, but above suspicion.
A shade of suspicion is often attached to dealers’ eggs \ not neces-
sarily implying bad faith or even negligence on the dealer’s part, but
from the nature of the case. It is often extremely difficult to make
an unquestionable determination, as, for instance, when numbers of
birds of similar habits are breeding close together ; or even impossible,
as in case the parent eludes observation. Sometimes the most acute
observer may be mistaken, circumstances appearing to prove a
parentage when such is not the fact. It is in general advisable to
secure the parent with the eggs ; if shot or snared on the nest, the
identification is unquestionable. If you do not yourself know the
species, it then becomes necessary to secure the specimen, and retain
it with the eggs. It is not required to make a perfect preparation ;
the head, or better, the head and a wing, will answer the purpose.
When egging in downright earnest, a pair of climbing irons, a coil
of f inch rope, and a tin collecting box filled with cotton, become
indispensable ; these are the only field implements required in
addition to those already specified.

Preparing’ Eggs. — For blowing eggs, a set of special tools is
needed. These are egg-drills, — steel implements with a sharp-
pointed conical head of rasping surface, and a slender shaft ; several
such, of different sizes, are needed ; also, blow- pipes of different
sizes, a delicate thin pair of scissors, light spring forceps, some little
hooks, and a small syringe. They are inexpensive, and may be had
of any dealer in naturalists’ supplies (see Figs. 4-7). Eggs^should
never be blowm in the old way of making a hole at each end ; nor
are two holes anywhere usually required. Opening should be
effected on one side, preferably that showing least conspicuous or

SEC. IX

COLLECTION OF NESTS AND EGGS

77

characteristic markings. If two are made, they should be rather
near together ; on the same side
at any rate. But one is generally
sufficient, as the fluid contents
can escape around the blow-pipe.

Holding the egg gently but
steadily in the Angers,^ apply the
point of the drill perpendicularly
to the surface, unless it be pre-
ferred to prick with a needle first.

A twirling motion of the instru-
ment gradually enlarges the open-
ing by filing away the shell, and
so bores a smooth-edged circular
hole. This should be no larger
than is required to insert the
blow-pipe loosely, with room for
the contents to escape around it.

Nor is it always necessary to
insert the pipe; a fine stream of
water may be easily injected by holding the instrument close to

the egg, but not
quite touching.
The blowing
should be contin-
uous and equable,
rather than for-
cible ; a strong
puff easily bursts
a delicate egg. Be
sure that all the
contents are re-
moved; then
rinse the interior
thoroughly with
clean water, either
by taking a
mouthful and
sending it
through a blow-
pipe, or with the
syringe. Blowing
eggs is a rather
fatiguing process ;

Fig. 5.— Instruments for blowing eggs ; after Newton, a, 6, blow-
pipes, i nat. size ; c, wire for cleansing them ; d, syringe, ^ nat. size
(the ring of the handle must be large enough to insert the thumb) ;
e, bulbous insufflator, for sucking eggs.

1 The usual method of emptying eggs through one small hole is doubtless

78

FIELD ORNITHOLOGY

PART I

the cheek -muscles soon tire, and the operator becomes “blown”
himself before long. The operation had better be done over
a basin of water, both to receive the contents, and to catch the
egg if it slip from the fingers. The membrane lining the shell

Fig. 6.— Scissors, knives, and forceps, J nat. size ;
after Newton.

should be removed if possible. It
may be seized by the edge around
the hole, with the forceps, and drawn
out, or picked out with a bent pin.

But this is scarcely to be accomp-
lished in the case of fresh eggs, when
the membrane may be simply pared
smoothly around the edge of the hole. Eggs that have been incubated
of course offer difficulty, in proportion to the size of the embryo. The

Fig. 7. — Hooks for extracting
embryos, nat. size ; after New-
ton, a, &, c, plain hooks ; d,
bill-hook, having cutting edge
along the concavity.

supposed to be a very modern trick ; but it dates back at least to 1828, when M.
Danger proposed “a new method of preparing and preserving eggs for the cabinet,”
which is practically the one now followed, though he used a three-edged needle to
prick the hole, instead of our modern drill, and did not appear to know some of our
ways of managing the embryo. I make this reference to his article to call attention
to one of the tools he recommends, which I think would prove useful, as being better
than the fingers for holding an egg during drilling and blowing. The simple instru-
ment will be understood from a glance at the figure given in the Nuttall Bulletin, iii.
1878, p. 191. The oval rings are covered with a light fabric, as mosquito-netting or
muslin, and do not touch the egg, which is held lightly but securely in the netting.
The cost would be trifling, and danger might be avoided by Danger’s method.

SEC. IX

COLLECTION OF NESTS AND EGGS

79

hole may be drilled, as before, but it must be larger ; and as the
drill is apt to split a shell after it has bored beyond a certain size
of hole, it is often well to prick, with a fine needle, a circular series
of minute holes almost touching, and then remove the enclosed
circle of shell. This must be very carefully done, or the needle will
indent or crack the shell, which, it must be remembered, grows
more brittle towards the time of hatching. Well-formed embryos
cannot be got bodily through any hole that can be made in an egg ;
they must be extracted piecemeal. They may be cut to pieces with
the slender scissors introduced through the hole, and the fragments
be picked out with the forceps, hooked out, or blown out. No
embryo should be forced through a hole too small j there is every
probability that the shell will burst at the critical moment. Addled
eggs, the contents of which are thickened or hardened, offer some
difficulty, to overcome which persistent syringing and repeated
rinsing are required ; or it may be necessary to fill them with water,
and set them away for such length of time that the contents dis-
solve by maceration ; carbonate of soda is said to hasten the solution \
the process may be repeated as often as may be necessary. In no event
must any of the animal contents be suffered to remain in the shell.
When emptied and rinsed, eggs should be gently wiped dry, and set
hole downward on blotting-paper to drain.^ Broken eggs may be neatly
mended, sometimes with a film of collodion, or a bit of tissue paper
and paste, or the edges may be simply stuck together with any
adhesive substance. Even when fragmentary a rare egg is worth
preserving. Eggs should ordinarily be left empty ; indeed, the
only case in which any filling is admissible is that of a defective

1 Reinforcing the Eggshell before Blowing. — Fig. 8 “ shows a piece of paper, a
Bumher of which, when gummed on to an egg, one over the other, and left to dry,
strengthen the shell in such a manner that the instruments above described can be
introduced through the aperture in the middle and worked to the best advantage, and
thus a fully formed embryo may be cut up, and the pieces
extracted, through a very moderately sized hole , the number
of thicknesses required depends, of course, greatly upon the
size of the egg, the length of time it has been incubated,
and the stoutness of the shell and the paper. Five or six
is the least number that it is safe to use. Each piece
should be left to dry before the next is gummed on. The
slits in the margin cause them to set pretty smoothly, which
will be found very desirable ; the aperture in the middle of
each may be cut out first, or the whole series of layers may
be drilled through when the hole is made in the egg. For
convenience’ sake, the papers may be prepared already
gummed, and moistened when put on (in the same way
that adhesive postage labels are used). Doubtless, patches of linen or cotton
cloth would answer equally well. When the operation is over, a slight application
of water (especially if warm) through the syringe will loosen them so that they can
be easily removed, and they can be separated from one another, and dried to serve
another time. The size represented in the sketch is that suitable for an egg of moder-
ate dimension, such as that of a common fowl. The most effectual way of adopting

Fig. 8.— Nat. size.

8o

FIELD ORNITHOLOGY

PART I

specimen to which some slight solidity can be imparted with cotton.
It is unnecessary even to close up the hole. It is best, on all
accounts, to keep eggs in sets^ a set being the natural clutch, or
whatever less number was taken from a nest. The most scrupu-
lous attention must be paid to accurate, complete, and permanent
labelling. So important is this, that the undeniable defacing of a
specimen, by writing on it, is no offset to the advantages accruing
from such fixity of record. It is practically impossible to attach a
label, as is done with a birdskin, and a loose label is always in
danger of being lost or displaced. Write on the shell, then, as
many items as possible ; if done neatly, on the side in which the
hole was bored, at least one good “ show side ” remains. An egg
should always bear the same number as the parent, in the collector’s
record. In a general collection, where separate ornithological
and oological registers are kept, identification of egg with parent is
nevertheless readily secured, by making one the numerator the
other the denominator of a fraction, to be simply inverted in its
respective application. Thus, bird No. 456 and egg No. 123 are
identified by making the former the latter -iff. All the eggs
of a clutch should have the same number. If the shell be large
enough, the name of the species should be written on it; if too
small, it should be accompanied by a label, and may have the name
indicated by a number referring to a certain catalogue. According
to my “ Check List,” for example. No. 4 would indicate Turdus
illacus, the common redwing. The date of collection is a highly
desirable item; it may be abbreviated thus: 3/6/82 means 3d June
1882. It is well to have the egg authenticated by the collector’s
initials at least. Since sets of eggs may be broken up for dis-
tribution to other cabinets, yet permanent indication of the size of
the clutch be wanted, it is well to have some method. A good one
is to write the number of the clutch on each egg composing it,
giving each egg of the set, moreover, its individual number. Sup-
posing, for example, the clutch No. contained five eggs ; one of
of them would be /5/1 : the next /5/2, and so on. But
it should be remembered that all such arbitrary memoranda must
be systematic, and be accompanied by a key. Eggs may be kept
in cabinets of shallow drawers in little pasteboard trays, each hold-
ing a set, and containing a paper label on which various items that

this method of emptying eggs is by using very many layers of thin 'paper and plenty
of thick gum, but this is, of course, the most tedious. Nevertheless, it is quite
worth the trouble in the case of really rare specimens, and they will be none the
worse for operating upon from the delay of a few days caused by waiting for the gum
to dry and harden. The naturalist to whom this method first occurred has found it
answer remarkably well in every case in which it has been used, from the egg of an
eagle to that of a humming-bird, and among English oologists it has been generally
adopted” (A. Newton, in Smithsonian Misc. Coll., p, 139, 1860).

SEC. IX

COLLECTION OF NESTS AND EGGS

8i

cannot be traced on the shell are written in full. Such trays should
all be of the same depth, — half an inch is a convenient depth for
general purposes ; and of assorted sizes, say from one inch by one
and one-half inches up to three by six inches j it is convenient to
have the dimensions regularly graduated by a constant factor of,
say half an inch, so that the little boxes may be set side by side, either
lengthwise or crosswise, without interference. Eggs may also be
kept safely, advantageously, and with attractive effect, in the nests
themselves, in which a fluff of cotton may be placed to steady them.
When not too bulky, too loosely constructed, or of material unsuit-
able for preservation, nests should always be collected.^ Those
that are very closely attached to twigs should not be torn off.
Nests threatening to come to pieces, or too frail to be handled with-
out injury, may be secured by sewing through and through with

1 “ A Plea for the Study of Nests,” made by Mr. Ernest Ingersoll in his excellent
Birds' -Nesting, suits me so well that I will transcribe it. “Whether or not it is
worth while to collect nests — for there are many persons who never do so — is, it
seems to me, only a question of room in the cabinet. As a scientific study there is
far more advantage to be obtained from a series of nests than from a series of eggs.
The nest is something with which the will and energies of the bird are concerned. It
expresses the character of the workman ; is to a certain extent an index of its rank
among birds, — for in general those of the highest organisation are the best architects,
— and give us a glimpse of the bird’s mind and power to understand and adapt itself
to changed conditions of life. Over the shape and ornamentation of an egg the bird
has no control, being no more able to govern the matter than it can the growth of its
beak. There is as much difference to me, in the interest inspired, between the nest
and the egg of a bird, as between its brain and its skull, — using the word brain to
mean the seat of intellect. The nest is always more or less the result of conscious
planning and intelligent work, even though it does follow a hereditary habit in its
style ; while the egg is an automatic production varying, if at all, only as the whole
organisation of the bird undergoes change. Don’t neglect the nests then. In them
more than anywhere else lies the key to the mind and thoughts of a bird, — the spirit
which inhabits that beautiful frame and bubbles out of that golden mouth. And is
it not this inner life, — this human significance in bird nature, — this soul of ornitho-
logy, that we are all aiming to discover ? Nests are beautiful, too. What can sur-
pass the delicacy of the humming-bird’s home glued to the surface of a mossy branch
or nestling in the warped point of a pendent leaf ; the vireo’s silken hammock ; the
oriole’s gracefully swaying purse ; the blackbird’s model basket in the flags ; the snug
little caves of the marsh wrens ; the hermitage-huts of the shy wagtails and ground
warblers, the stout fortresses of the sociable swallows ! Moreover, there is much that
is highly interesting which remains to be learned about nests, and which can only be
known by paying close attention to these artistic masterpieces of animal art. We
want to know by what sort of skill the many nests are woven together that we find
it so hard even to disentangle ; we want to know how long they are in being built ;
whether there is any particular choice in respect to location ; whether it be a rule,
as is supposed, that the female bird is the architect, to the exclusion of her mate’s
efforts further than his supplying a part of the materials. Many such points remain to
be cleared up. Then there is the question of variation, and its extent in the archi-
tect of the same species in different quarters of its ranging area. How far is this
carried, and how many varieties can be recorded from a single district, where the same
list of materials is open to all the birds equally ? Variation shows individual opinion
or taste among the builders as to the suitability of this or that sort of timber or fur-
niture for their dwellings, and observations upon it thus increase our acquaintance
with the scope of ideas and habits characteristic of each species of bird.”

G

82

FIELD ORNITHOLOGY

PART I

fine thread : indeed, this is an advisable precaution in most cases.
Packing eggs for transportation requires much care, but the pre-
cautions to be taken are obvious. I will only remark that there is
no safer way than to leave them in their own nests, each wrapped
in cotton, with which the whole cavity is to be lightly filled ; the
nests themselves being packed close enough to be perfectly steady.

§ 10.— CAEE OF A COLLECTION

Well-preserved Specimens will last “ for ever and a day,” so
far as natural decay is concerned. I have handled birds in good
state, shot back in the twenties, and have no doubt that some
eighteenth-century preparations are still extant. The precautions
against defilement, mutilation, or other mechanical injury, are self-
evident, and may be dismissed with the remark, that white plum-
ages, especially if at all greasy, require the most care to guard
against soiling. We have, however, to fight for our possessions
against a host of enemies, individually despicable but collectively
formidable, — foes so determined that untiring vigilance is required
to ward off their attacks even temporarily, whilst in the end they
prove invincible. It may be said that to be eaten up by insects is
the natural end of all bird-skins not sooner destroyed.

Insect Pests (Figs. 9, 10, 11, 12) with which we have to con-
tend belong principally to the two families Tineidce and Dermestidce
— the former are moths, the latter beetles. The moths are of
species identical with, and allied to, the common clothes-moth.
Tinea flavifrontella, the carpet moth, T. ta;petzella, etc., — small species
observed flying about our apartments and museums, in May and
during the summer. The beetles are several lather small thick-set
species, principally of the genera Dermestes and Anthrenus. I am
able to figure species of these genera, with their larval stages, and
of two other genera, Ptinus and Sitodrepa, through the attentions of
Professor C. V. Riley, the eminent entomologist. The larvae (“ cater-
pillars ” of the moths, and “ grubs ” of the beetles) appear to be
the chief agents of the destruction. The presence of the mature
insects is usually readily detected ; on disturbing an infested suite
of specimens the moths flutter about, and the beetles crawl as fast
as they can into shelter, or simulate death. The insidious larvae,
however, are not so easily observed, burrowing as they do among
the feathers, or in the interior of a skin ; whilst the minute eggs
are commonly altogether overlooked. But these insects are not
long at work without leaving their unmistakable traces. Shreds
of feathers float off when a specimen is handled, or fly out on flip-

SEC. X

CA/^E OF A COLLECTION

83

ping the skin with the fingers, and in bad cases even whole bundles
of plumes come away at a touch. Sometimes, leaving the plumage
intact, insects eat away the horny covering of the bill and feet,
making an irreparable mutilation. It would appear that when the
pests effect lodgment in any one skin, they usually finish it before
attacking another, unless they are in great force. We may con-

Fig. 9. — AnfhreniLs scroplmlari^, enlarged ; the short line shows nat. size, a, h, larvae ;
c, pupa; cl, imago.

•Fig. 10. — Dermestes lardarius, Fig. 11. — Sitodrepa panicea, Fig. 12.— Ptinus hmnneus.

enlarged, a, larva ; I, an enlarged enlarged, a, imago ; b, its an-
hair ; c, imago. tenna, more enlarged.

sequently, by prompt removal of an infested specimen, save further
depredations ; nevertheless, the rest become suspicious, and the
whole drawer or box should be quarantined, if not submitted to
any of the processes described beyond. Our lines of defence are
several. We may mechanically oppose entrance of the enemy; we
may meet him with abhorrent odours that drive him off*, sicken or
kill him, and finally we may cook him to death. I will notice
these methods successively, taking occasion to describe a cabinet
under the first head.

84

FIELD ORNITHOLOGY

PART I

Cases fop Storage op Transpoptation should be rather small,
for several reasons. They are easier to handle and pack. There
are fewer birds pressing each other. Particular specimens are more
readily reached. Insects must effect just so many more separate
entrances to infest the whole. Small lids are more readily fitted
tight. For the ordinary run of small birds I should not desire a
box over 18x18x18 inches, and should prefer a smaller one ; for
large birds, a box just long enough for the biggest specimen, and of
other proportions to correspond, is most eligible. Whatever the
dimensions, a proper box presupposes perfect jointing ; but if any
suspicion be entertained on this score, stout paper should be pasted
along all the edges, both inside and out. We have practically to
do with the lid only. If the lot is likely to remain long untouched,
the cover may be screwed very close and the crack pasted like the
others. Under other and usual circumstances the lid may be pro-
vided with a metal boss fitting a groove lined with india-rubber or
filled with wax. An excellent case may be made of tin with the
lid secured in this manner, and further fortified with a wooden
casing. Birdskins entirely free from insects or their eggs, encased
in some such secure manner, will remain intact indefinitely; but
there is misery in store if any bugs or nits be put away with them.

Cabinets. — As a matter of fact, most collections are kept readily
accessible for examination, display, or other immediate use, and this
precludes any disposition of them in hermetical cases. The most
we can do is to secure tight fitting of movable woodwork. The
cabinet is most eligible for private collections. This is, in effect,
simply a bureau, or chest of drawers, protected with folding doors,
or a front that may be detached, either of plain wood or sashing
for panes of glass. It is astonishing how many birdskins of average
size can be accommodated in a cabinet that makes no inconvenient
piece of furniture for an ordinary room. A cabinet may of course
be of any desired size, shape, and style. In general it will be
better to put money into excellence of fitting rather than elegance
of finish ; the handsomest front does not compensate for a crack in
the back or for a drawer that hitches. There should not be the
slightest fiaw in the exterior, and doors should fit so tightly that a
puff of air may be felt on closing them. The greatest desideratum
of the interior work, next after close fitting yet smooth running of
the drawers, is economy of space. This is secured by making the
drawers as thin as is consistent with stability ; by having them
slide by a boss at each end fitting a groove in the side wall, instead
of resting on horizontal partitions ; and by hinged countersunk
handles instead of knobs. I do not recommend, except for a suite
of the smallest birds, a multiplicity of shallow drawers, accommodat-
ing each one layer of specimens ; it is better to have fewer deeper

SEC. X

CAJ?E OF A COLLECTION

85

drawers, into which light shallow movable trays are fitted. These
trays never need be of stuff over one-eighth or one-fourth of an
inch thick, and may have bottoms of stiff pasteboard glued or
tacked on. They may vary from one-half inch to two inches in
depth, but this dimension should always be some factor of the
depth of the drawer, so that a certain number of trays may exactly
fill it. They should be just as long as one transverse dimension of
the drawer, and rather narrow, so that two or more are set side by
side. Finally, though they may be of different depths, they should
be of the same length and breadth, so as to be interchangeable.
They may simply rest on top of each other, or slide on separate
projections inside the drawer. Such trays are extremely handy
for holding particular sets of specimens, to be carried to the study
table without disturbing the rest of the collection.

If a collection be so extensive that any particular specimen may
not be readily hunted up, it will be found convenient to have the
drawers themselves labelled with the name of the group within.
A collection should always be methodically arranged — preferably
according to some approved or supposed natural classification of
birds j this is also the readiest mode, since, with some conspicuous
exceptions, birds of the same natural group are approximately of
the same size. If I were desired to suggest proportions for a private
cabinet of most general eligibility, I should say four feet high, by
three feet wide, by two feet deep, in the clear ; this makes a portly
yet not unwieldy-looking object. It is wide enough for folding-
doors, to be secured by bolts at top and bottom, and lock ; not so
high that the top drawer is not readily inspected ; and of propor-
tionate depth. Such a case will take seven drawers six inches deep
either of the full width or in two series with a median partition ;
these drawers will hold anything up to an eagle or crane. A part
of them at least should have a full complement of such trays as I
have described, — say three or four tiers of the shallower trays, three
trays to a tier, each about two feet long by about a foot wide ; and
one or two tiers of deeper trays.

To Destroy Insects. — In our present case prevention is not the
best remedy, simply because it is not always practicable ; in spite
of all mechanical precautions the insects will get in. We have,
therefore, to see what will destroy them, or at least stop their
ravages. It is a general rule that any pungent aromatic odour is
obnoxious to them, and that any very light powdery substance
restrains their movements by getting into the joints and breathing
pores. Both these qualities are secured in the ordinary insect
powder, to be had of any leading druggist. It should be lavishly
strewn on and among the skins, and laid in the corners of the
drawers and trays. Thus employed it proves highly effective, and

86

FIELD ORNITHOLOGY

PART I

is on the whole the most eligible substance to use when a collection
is constantly handled. Camphor is a valuable agent. Small
fragments may be strewn about the drawers, or a lump pinned in
mosquito netting in a corner. Benzine is also very useful. A
small saucerful may be kept evaporating, or the liquid may be
sprinkled — even poured — directly over the skins ; it is very volatile
and leaves little or no stain. It is, however, obviously ineligible
when a collection is in constant use. My friend Mr. Allen informs
me he has used sulphide of carbon with great success. The
objection to this agent is, that it is a stinking poison ; should be
used in the open air, to escape the ineffably disgusting and dele-
terious odours, and its employment is properly restricted to cases for
storage. When the bill or feet show they are attacked, further
depredation may be prevented by pencilling with a strong solution
of corrosive sublimate ; a weaker solution, one that leaves no white
film, on drying, on a black feather, may even be brushed over the
whole plumage. Mr. Kidgway tells me that oil of bitter almonds
is equally efficacious. But remember that these poisons must be
used with care. Specimens may be buried in coarse refuse tobacco
leaves. One or another of these lines of defence will commonly
prove successful in destroying or driving off mature insects, and
even in stopping the ravages of the larvae j but I doubt that any
such means will kill the eggs. With these w^e must deal otherwise ;
and their destruction no less than that of their parents is assured,
if we subject them to a high temperature. Baking birdskins is
really the only process that can make us feel perfectly safe.
Infected specimens, along with suspected ones, should be subjected
to a dry heat, from 212° F. up to any degree short of singeing the
plumage. This is readily done by putting the birds in a wooden
tray in any oven — they must however be watched, unless you have
special contrivances for regulating the temperature. How long a
time is required is probably not ascertained with precision ; it will
be well to bake for several hours. When the beetles and larvae are
found completely parched, it may be confidently believed that the
unseen eggs are out of the hatching way for ever.

Arsenic helps to keep out the bugs, besides preventing decay — a
fact that should never be forgotten, and that should give sharper edge
to my advice respecting lavish use of the substance at the outset. If
it be true, as some state, that bugs can eat arsenic without dying,
it is also true that they do not relish it ; and in entering a case of
skins they will burrow by preference in those holding the least of it.
This fact is continually exhibited in large collections, where if two
birds be side by side, one being duly arsenicised and the other not
so, one will be taken and the other left. It is also a fact in the
natural history of these our pests, that they are fond of peace and

SEC. X

CAJ?E OF A COLLECTION

87

quiet, — they do not like to be disturbed at their meals. So they
rarely effect permanent lodgment in a collection that is constantly
handled, though the doors stand open for hours daily. As a con-
sequence, the degree of our diligence in studying birdskins is likely
to become the measure of our success in preserving them. I once
read a work, by an eminent divine, on the Moral Uses of Dark
Things, under which head the author included every dark thing from
earthquakes to mosquitoes. If there be a moral use in the “ dark
thing” that museum pests certainly are to us, we have it here.
The very bugs urge on our work.

Fig. 13. — Alexander Wilson’s School-house, near Gray’s Ferry, Philadelphia, U.S.A.
From a drawing by M. S. Weaver, Oct. 22, 1841, received by Elliot Cones, February 1879, from
Malvina Lawson, daughter of Alexander Lawson, Wilson’s engraver. See article in the Penn
Monthly, June 1879, p. 443. The drawing was first engraved on wood, and published, by
Thomas Meehan, in the Gardener’s Monthly, August 1880, p. 248. The present impression is
from an electrotype of that woodcut. The size of the original is 5T0x3‘95 inches. This re-
minder of early days of “Field Ornithology” in America maybe further attested by the
signature of

PAET II

GENEEAL OENITHOLOGY

AN OUTLINE OF THE

STKUCTUKE AND CLASSIFICATION OF BIEDS

GENEEAL OENITHOLOGY

§ 1.— definition of bieds

General Ornithology, like Field Ornithology, is a subject with
which the student must have some acquaintance, if he would hope
to derive either pleasure or profit from the Birds of Great Britain.
For any intelligent understanding of this subject, he must become
reasonably familiar with the technical terms used in describing and
classifjdng birds, and learn at least enough of the structure of these
creatures to appreciate the characters upon which all description
and classification is based. Extensive and varied and accurate as
may be his random perception of objects of natural history, his
knowledge is not scientific, but only empirical, until reflection comes
to aid observation, and conceptions of the significance of what he
knows are formed by logical processes in the mind. For

Science (Lat. scire, to know) is knowledge set in order ; know-
ledge disposed after the rational method that best shows, or
tends to show best, the true relations of observed facts. Sound
scientific facts are the natural basis of all philosophic truth, and
the safest stepping-stones to religious faith, — to that wisdom
which comes only of knowing the relation which material
entities bear to spiritual realities. The orderly knowledge of
any particular class of facts — the methodical disposition of observa-
tions upon any particular set of objects — constitutes a Special
Science. Thus

Ornithology (Gr. opvtOos, ornithos, of a bird ; Aoyo?, logos, a
discourse) is the Science of Birds. Ornithology consists in the
rational arrangement and exposition of all that is known of birds,
and the logical inference of much that is not known. Ornithology
treats of the physical structure, physiological processes, and mental
attributes of birds ; of their habits and manners ; of their geo-

92

GENERAL ORNITHOLOGY

PART II

graphical distribution and geological succession ; of their probable
ancestry ; of their every relation to one another and to all other
animals, including man, — in short, of their significance in Nature.
The first business of Ornithology is to define its ground — to answer
the question

What is a Bird ? — There is every reason to believe that a Bird
is a greatly modified Eeptile, being the offspring by direct descent
of some reptilian progenitor; and there is no reason to suppose
that any bird ever had any other origin than by due process of
hatching out of an egg laid by its mother after fecundation by its
father, — just what we believe to have been the invariable method
during the period of the world known to human history. There is
no reason to believe that any bird was ever originally created and
endowed with the characters it now possesses ; but that every bird
now living is the naturally modified lineal descendant of parents
that were less and less like itself, and more and more like certain
reptiles, the further removed they were in the line of avian
ancestry from such birds as are now living. This is the Darwinian
logic of observed facts, upon which the modern Theory of Evolution
is based, in opposition to the tradition of the special creation of
every species of animal ; which latter has no scientific basis
whatever, and is consequently accepted as true by few thoughtful
persons who are capable of forming independent judgments.
Accordingly,

Birds and Reptiles — even those of the present geologic epoch
— share so many and so important structural characters, that the
chiefs of science of our day are wont to unite the two classes, Ams
and Reptilia, in one primary group of the Vertehrata, or animals
with a backbone. This group is called Sauropsida, or reptiliforms ;
it is contrasted, on the one hand, with Ichthyopsida, or fish -like
vertebrates, including Batrachians as well as Fishes ; and, on the
other, with Mammalia, the province of the Vertebrata which in-
cludes Man and all other animals that suckle their young. We
find that

The Sauropsida (Gr. o-avpos, sauros, a reptile ; 6\pLs, opsis, appear-
ance), or lizard-like Vertebrates, agree with one another, and differ
from other animals, in the following important combination of char-
acters, substantially as laid down by Professor Huxley, — some of
the characters being shared by the Ichthyopsida, and some by the
Mammalia, but the sum of the characters being distinctive of Saurop-
sida : they are all oviparous (laying eggs hatched outside the body
of the parent), or ovoviviparous (laying eggs hatched inside the body
of the parent), being never viviparous (bringing forth alive young
nourished before birth by the blood of the mother). The embryo
develops those foetal organs called amnion and allantois, and is

SEC. I

DEFINITION OF BIRDS

93

nourished before hatching by the great quantity of food-yolk in the
egg. There are no mammary glands to furnish the young with
milk after birth. The generative, urinary, and digestive organs
come together behind in a common receptacle, the cloaca, or sewer,
and their products are discharged by a single orifice. The kidneys
of the early embryo, called Wolffian bodies, soon replaced func-
tionally by permanent kidneys, and structurally by the testes of the
male and the ovaries of the female. The cavity of the abdomen, or
belly, is not separated from that of the tlmax, or chest, by a com-
plete muscular partition, or diaffiiragm. The great lateral hemi-
spheres of the brain are not connected by a transverse commissure,
or corpus callosum. Air is always breathed by true lungs, never by
gills. The blood, which may be cold or hot, has red oval nucleated
corpuscles ; the heart has either three or four separate chambers, —
the latter in birds, in which the circulation of the hot blood is com-
pletely double, i.e. in the lungs and one side of the heart, in the
body at large and the other side of the heart. The aortic arches are
several ; or if but one, as in birds, it is the right, not the left as in
mammals. The centra, or bodies, of the vertebrae are ossified, but
have no terminal epiphyses. The skull hinges upon the backbone by
a single median protuberance, or condyle, and the bone {basioccipital)
bearing this condyle is completely ossified. The lower jaw consists of
several separate pieces, the articular one of which hinges upon a mov-
able quadrate bone ; and there are other peculiarities in the formation
of the skull. The ankle-joint is situated, not, as in Mammals, between
the tarsal bones and those of the leg, but between two rows of tarsal
bones. The skin is usually covered with outgrowths, in the form of
scales or feathers. Different as are any living members of the class
of Birds from any known Eeptiles, the characters of the two groups
converge in geologic history so closely, that the presence oi feathers
in the former class, and their absence from the latter, is one of the
most positive differences we have found. The oldest known birds
are from the Jurassic rocks of Europe, and the Cretaceous beds of
North America. These birds had teeth, and various other strong
peculiarities of structure, which no living members of the class have
retained.

AVES, OP the Class of Birds, may be distinguished from other
Sauropsida, for all that is known to the contrary, by the following
sum of characters : The body is covered with feathers, a kind of
skin- outgrowth no other animals possess. The blood is hot; the
circulation is completely double ; the heart is perfectly four-
chambered ; there is but one (the right) aortic arch, and only one
pulmonary artery springs from the heart ; the aortic and the pul-
monary artery have each three semilunar valves. The lungs are
fixed and moulded to the cavity of the chest, and some of the air-

94

GENERAL ORNITHOLOGY

PART II

passages run through them to admit air to other parts of the body,
as under the skin and in various bones. Eeproduction is oviparous ;
the eggs are very large, in consequence of their copious yolk and
white \ have a hard chalky shell, and are hatched outside the body
of the parent. There are always four limbs, of which the fore
or pectoral pair are strongly distinguished from the hind or pelvic
pair by being modified into wings^ fitted for flying, if at all, by
means of feathers — not of skin as in the cases of such mammals,
reptiles, and fishes as can fly. The terminal part of the limb is
compressed and reduced, bearing never more than three digits, only
two of which ever have claws, and no claws being the rule. There
are not more than two separate car'pals, or wrist-bones, in adult
recent birds (with very rare exceptions) ; nor any distinct inter-
clavicular bone. The clavicles are complete (with rare exceptions),
and coalesce to form a “wish-bone” or “merrythought.” The
sternum, or breast-bone, is large, usually carinate, or keeled, and the
ribs are attached to its sides only ; it is developed from two to five
or more centres of ossification. The sacral vertebrae proper have no
expanded ribs abutting against the ilia ; the ilia, or haunch-bones,
are greatly prolonged forward ; the socket for the head of the
femur, or thigh-bone, is a ring, not a cup ; the ischia and pules are
prolonged backward in parallel directions, and neither of these bones
ever unites with its fellow in a ventral symphysis (except in Struthio
and Rhea). The fibula, or outer bone of the leg, is incomplete
below, taking no part in the ankle-joint. The astragalus, or upper
bone of the tarsus, unites with the tibia, or inner bone of the leg,
leaving the ankle-joint between itself and other tarsal bones, the
lower of which latter similarly unites with the bones of the instep,
or metatarsus. There are never more than four metatarsal bones,
and the same number of digits ; the first or inner metatarsal bone
is usually free, and incomplete above ; the other three anchylose
(fuse) together, and with distal tarsal bones, as already said, to form
a compound tarso-metatarsus. Eecent birds, at any rate, have a
certain saddle-shape of the ends of the bodies of some vertebrae.
Such birds have also no teeth and no fleshy lips ; the jaws are
covered with horny or leathery integument, as the feet are also,
when not feathered.

The Position of the Class Aves among other Vertebrates is
definite. Birds come in the scale of development next below the
Class Mammalia, and no close links between Birds and Mammals are
known ; the most bird-like known mammal, the duck-billed platypus
of Australia {Ornithorhynchus paradoxus), being several steps beyond
any known bird. Birds are the higher one of the two classes of
Sauropsida — the lower class, Reptilia, connecting with the Batrachians
(frogs, toads, newts, etc.), and so with the Fishes, IcMhyopsida. In

SEC. I

DEFINITION OF BIRDS

95

this Vertebrate series, Birds constitute what is called a highly
s;pecialised group ; that is to say, a very particular offshoot, or, more
literally, a side-issue, of the Vertebrate genealogical tree, which in
the present geological era has become developed into very numerous
(about 10,000) species, closely agreeing with one another in the
peculiar sum of their physical character. In comparison with other
classes of Vertebrates, all birds are much alike ; there is a less degree
of difference among them than that found among the members of
any of the other classes of Vertebrates ; their likeness to each other
being strong, and their kind of difference from any other Verte-
brates being peculiar, makes them the highly specialised class
they are recognised to be. The structural difference between a
humming-bird and an ostrich, for example, is not greater in degree
than that subsisting between the members of some of the orders of
Eeptiles ; whence some hold, with reason, that Birds should not form
a class Aves, but an order, or at most a sub-class, of Sauropsida, and
so be compared not with a class Beptilia collectively, but with other
sauropsidan orders, such as Chelonia (turtles), Sauria (lizards),
Ophidia (serpents), etc. The practical convenience of starting with
a “class” Aves, however, is so great, that such classificatory value
will probably long continue to be ascribed, as heretofore, to Birds
collectively. I have spoken of Birds as a particular side-issue or
lateral branch of the Vertebrate “ tree of life ” ; hence it is not to be
supposed that they are in the direct line of genealogical descent.
Though they stand as a group next below Mammals in the scale of
evolution, it does not follow that Mammals were developed from
any such creature as a Bird has come to be, any more than that
Birds have been evolved from any such Eeptiles as those of the pre-
sent day. It is one of the popular misunderstandings of the Theory
of Evolution, to imagine that all the lower forms of animals are in
the genetic line of development of the higher forms ; that man, for
example, was once a gorilla or a chimpanzee — actually such an ape.
The theory simply requires all forms of life to be developed from
some antecedent form, presumably, and in most cases certainly, lower
in the scale of organisation. Thus man and the gorilla are both
descendants of some common progenitor, more or less unlike either
of these existing creatures. All mammals are similarly the modified
descendants of some more primitive stock, from which stock sprang
also all Sauropsida, mediately or immediately ; therefore a Mammal
is not a modified Bird, though higher in the scale ; and, though a
Bird is a modified Eeptile, it is not a modification of any such
snake or lizard as now exists. The most bird-like reptiles known
are not the Pterodactyls, or Flying Eeptiles (Pterosmiria), as might
be supposed ; but belong to that remarkable order, the Ornitho-
scelida, comprising the Dinosaurians, which “ present a large series of

96

GENERAL ORNITHOLOGY

PART II

modifications intermediate in structure between existing
and Aves^'' and are therefore inferentially in the direct ancestral
line of modern Birds.

Geologic Succession of Binds. — Birds have been traced back in
geologic time to Cretaceous and Jurassic epochs of the Mesozoic or
Mid-Life period of the world’s history. The earliest ornithichnites, —
the fossils so called because supposed to indicate the presence of

Birds by their
footprints, were
discovered about
the year 1835 in
the Triassic for-
mation in Con-
necticut. But the
creatures which
made these tracks
are now reason-
ably believed to
have been all
Dinosaurian rep-
tiles. The oldest
ornithoUte, or fos-
sil certainly
known to be that
of a true Bird, is
the famous
o;pteryx, found by
Andreas Wagner
in 1861 in the
Oolitic slate of
Solenhofen in
Bavaria. This has
a long lizard-like
tail of twenty ver-
tebrae, from each
of which springs
feathers of the wings

Fig. 14.— Oldest known ornithological treatise, illustrating also
the art of lithography in the Jurassic period, engraved by ArcTuBO-
pteryx lithograpMca. From the original slab in the British Museum ;
after A. Newton, Ency. Brit.

a well-developed feather on each side ; ^

are also well preserved; bones of the hand are not fused to-
gether, as they are in recent Birds; and the jaws hear true
teeth ’ This Bird has served as the basis of one of the primary divi-
sions of the class Aves; though it has many reptilian characters, it
is a true Bird. The great gap between this ancient Avian and
latter-day birds has been to some extent bridged by the discovery
and restoration of Birds from the Cretaceous formations of North
America such genera as Ichthyornis and Hesperornis forming types of

SEC. I

DEFINITION OF BIRDS

97

two other primary divisions of the class, Odontotormm and Odontolcce,
or Birds with teeth in sockets, and those with teeth in grooves. In
both these genera the tail is short, as in ordinary birds. In Iclithy-
ornis, though the wings are well developed, with fused metacarpals,

Fig. 15.— Restoration of Hesperornis regalis. After Marsh.

and the sternum is keeled, the vertebrae present the extraordinary
primitive character of being biconcave. In Hesperornis the vertebrae
are saddle-shaped, as usual, but the sternum is flat, as in the existing
ostriches, and the wings are rudimentary, wanting metacarpals.
Some twenty species of several genera of other American Cretaceous
H

98

GENERAL ORNITHOLOGY

PART II

Birds have been described. Eemains of Birds multiply in
the next period, the Tertiary. Those of the Eocene or early
Tertiary are largely and longest known from discoveries made
in the Paris Basin, among them the Gastornis jparisiensis^ at

Fig. 16. — Restoration of Ichthyornis victor. After Marsh.

least as large as an ostrich ; some of these belong to extinct
genera, others to genera which still flourish ; none are known
to have true teeth, or otherwise to be as primitive as the reptile-like
forms of the Cretaceous. The Miocene or Middle Tertiary has
proved specially rich in remains of Birds, including some of extinct

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 99

genera, but in largest proportion referable to modern types. Later
Tertiary (Pliocene and Post-pliocene) birds are almost all of living
genera, and some are apparently of living species. Extinct birds
coeval with man, their bones bearing his
marks, are found in various caves. Sub-fossil
birds’ bones occur in shell -heaps (kitchen-
middens) and elsewhere, of course contempor-
aneous with man, and some of them scarcely
prehistoric. One of the oldest of these is the
gigantic j^pyornis maximus of Madagascar, of
which we have not only the bones, but the
egg. The immense Moas, or Dinornithes of
New Zealand, were among the later of these
to die, portions of skin, feathers, etc., of
these great creatures having been found.

With the Moa-remains are found those of
Harpagornis^ a raptorial bird large enough to
have preyed upon the Moas. Finally, various
birds have been exterminated in historic
times, and some of them within the lifetime
of persons now living. The Dodo of Mauri-
tius, Didusineptus^ is the most celebrated one
of these, of the living of which we have
documentary evidence down to 1681 ; the ■

Solitaire of Eodriguez, Fezophaps solitarius,
the Geant, Leguatia gigantea, and several
others of the same Mascarene group of islands,
are in similar case. The Great Auk, Aka LegiiAla'
impenniSj is supposed to have become extinct

in 1844; a species of Parrot, Nestor productus, was last known to
be living in 1851 ; various parrots and other birds have likewise
disappeared within a very few years. At least one North American
bird, the Labrador Duck, Camptolcemus lahradorius, seems likely
soon to follow. (A. Newton, Ency. Brit, ninth edition, art. ‘Birds.’)

§ 2.— PRINCIPLES AND PRACTICE OF CLASSIFICATION

Having seen what a Bird is, and how it is distinguished from
other animals, our next business is to inquire how birds are related
to and distinguished from one another, as the basis of

Classification : a prime object of ornithology, without the
attainment of which birds, however pleasing they are to the senses,
do not satisfy the mind, which always strives to make orderly

lOO

GENERAL ORNITHOLOGY

PART II

disposition of its knowledge, and so discover the reciprocal relations
and interdependencies of the things it knows. Classification pre-
supposes that there do exist such relations, according to which we
may arrange objects in the manner which facilitates their compre-
hension, by bringing together what is like, and separating what is
unlike j and that such relations are the results of fixed, inevitable
law. It is, therefore,

Taxonomy (Gr. rafi?, taxis, arrangement, and vo/xo?, nomos, law),
or the rational, lawful disposition of observed facts. Just as taxi-
dermy is the art of fixing a bird’s skin in a natural manner, so
taxonomy is the science of arranging birds in the most natural
manner; in the way that brings out most clearly their natural
affinities, and so shows them in their proper relations to one another.
This is the greatest possible help to the memory in its attempt to
retain its hold upon great numbers of facts. But taxonomy, which
involves consideration of the greatest problems of ornithology, as of
every other branch of biology (biology being the science of life and
living things in general), is beset with the gravest difficulties,
springing from our defective knowledge. We could only perfect
our taxonomy by having before us a specimen of every kind of bird
that exists, or ever existed ; and by thoroughly understanding how
each is related to and differs from every other one. This is obvi-
ously impossible ; in point of fact, we do not know all the birds
now living, and only a small number of extinct birds have come to
light ; so that many of the most important links in the chain of
evidence are missing, and many more cannot be satisfactorily joined
together. With these springs of ignorance and sources of error
must be reckoned also the risk of going wrong through the natural
fallibility of the mind. The result is, that the “natural classifica-
tion,” like the elixir of life or the philosopher’s stone, is a goal still
distant ; and as a matter of fact, the present state of the ornitho-
logical system is far from being satisfactory. It is obvious that
birds, or any other objects, may be classified in numberless ways,
— in as many ways as are afforded by all their qualities and rela-
tions,— to suit particular purposes, or to satisfy particular bents of
mind. Hence have arisen, in the history of the science, very many
different schedules of classification ; in fact, nearly every leader of
ornithology has in his time proposed his own system, and enjoyed
a more or less respectable and influential following. Systems have
been based upon this or that set of characters, and erected from
this or that preconception in the mind of the systematist. Down
to quite recent days, the modifications of the external parts of birds,
particularly of the bill, feet, wings, and tail, were almost exclusively
employed for purposes of classification; and the mental point of
view was, that each species of bird was a separate creation, and as

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION loi

much of a fixture in Nature’s museum as any specimen in the
naturalist’s cabinet. Crops of classifications have been sown in the
fruitful soil of such blind error, but no lasting harvest has been
reaped. The confusion thus engendered has brought about the
inevitable reaction ; and the fashion of the present day is decidedly
the opposite extreme, — that of counting external features of little
consequence in comparison with anatomical characters. Too much
time has been wasted in arguing the superiority of each of these
characters for the purposes of classification ; as if a natural classi-
fication should not be based upon all points of structure ! as if
internal and external characters were not reciprocal and mutually
exponent of each other ! But the genius of modern taxonomy
seems to be so certainly right, — to be tending so surely, even if
slowly, in the direction of the desired consummation, that all
differences of opinion, we may hope, soon will be settled, and
defect of knowledge, not perversity of the mind, be the only
obstacle left in the way of success. The taxonomic goal is not now
to find the way in which birds may be most conveniently arranged,
described, and catalogued ; but to discover their pedigree, and so
construct their family-tree. Such a genealogical table, or phylum
(Gr. (fivXov, pliulon, tribe, race, stock), as it is called, is rightly con-
sidered the only taxonomy worthy the name, — the only true or
natural classification. In attempting this end, we proceed upon
the belief that, as explained above, all birds, like all other animals
and plants, are related to each other genetically, as offspring are to
parents ; and that to discover their genetic relationships is to bring
out their true affinities, — in other words, to reconstruct the actual
taxonomy of Nature. In this view, there can be but one natural
classification, to the perfecting of which all increase in our knowledge
of the structure of birds infallibly tends. The classification now in
use, or coming into use, is the result of our best- endeavours to
accomplish this purpose, and represents what approach we have
made to this end. It is one of the great corollaries of that theorem
of Evolution which most naturalists are satisfied has been demon-
strated. It is necessarily a

Morphological Classification ; that is, one based solely upon
consideration of structure or form (fjbop4>'q, morphe, form) ; and for
the following reasons : Every offspring tends to take on precisely
the structure or form of its parents, as its natural physical heritage j
and the principle involved, or the law of heredity, would, if nothing
interfered, keep the descendants perfectly true to the physical
characters of their progenitors ; they would “ breed true ” and be
exactly alike. But counter-influences are incessantly operative, in
consequence of constantly varying external conditions of environ-
ment ; the plasticity of organisation of all creatures rendering them

102

GENERAL ORNITHOLOGY

PART II

more or less susceptible of modification by such means, they become
unlike their ancestors in various ways and to different degrees. On
a large scale is thus accomplished, by natural selection and other
natural agencies, just what man does in a small way in producing
and maintaining different breeds of domestic animals. Obviously,
amidst such ceaselessly shifting scenes, degrees of likeness or unlike-
ness of physical structure indicate with the greatest exactitude the
nearness or remoteness of organisms in kinship. Morphological
characters derived from examination of structure are therefore the
surest guides we can have to the blood-relationships we desire to
establish ; and such relationships are the natural affinities which
all classification aims to discover and formulate. As already said,
taxonomy consists in tracing pedigrees, and constructing the jpliylum ;
it is like tracing any leaf or twig of a tree to its branchlet, this to
its bough, this again to its trunk or main stem. The student will
readily perceive, from what has been said, the impossibility of
naturally arranging any considerable number of birds in any linear
series of groups, one after the other. To do so means nothing more
or less than the mechanical necessity of book-making, where groups
have to succeed one another, in writing page after page. Some
groups will follow naturally ; others will not ; no connected chain
is possible, because no such single continuous series exists in nature.
In cataloguing, or otherwise arranging a series of birds for descrip-
tion, we simply begin with the highest — or lowest, if we prefer —
groups, and make our juxtapositions as well as we can, in order to
have the fewest breaks in the series.

Morphology being the safest, indeed the only safe, clue to
natural affinities, and the key to all rational classification, the
student cannot too carefully consider what is meant by this term,
or too sedulously guard against misinterpreting morphological char-
acters, and so turning the key the wrong way. The chief difficulty
he will encounter comes from physiological adaptations of structure ;
and this is something that must be thoroughly understood. The
expression means that birds, or any animals, widely different in the
• sum of their morphological characters, may have certain parts of
their organisation modified in the same way, thus bringing about a
seemingly close resemblance between organisms really little related
to each other. For example : a phalarope, a coot, and a grebe, all
have lobate feet ; that is, their feet are fitted for swimming purposes
in the same way, namely, by development of flaps or lobes on the
toes. A striking but very superficial and therefore unimportant
resemblance in a certain particular exists between these birds, on
the strength of which they used to be classed together in a group
called Pinnatipedes, or “fin-footed” birds. But, on sufficient ex-
amination, these three birds are found to be very unlike in other

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 103

respects ; the sum of their unlikenesses requires us to separate
them quite widely in any natural system. The group Finnatipedes
is therefore unnatural, and the appearance of affinity is proved to
be deceptive. Such resemblance in the condition of the feet is
simply functional, or physiological, and is not correspondent with
structural or morphological relationships. The relation, in short,
between these three birds is analogical ; it is an inexact superficial
resemblance between things profoundly unlike, and therefore having
little homological or exact relationship. Analogy is the apparent
resemblance between things really unlike, — as the wing of a bird
and the wing of a butterfly, as the lungs of a bird and the gills of
a fish. Homology is the real resemblance or true relation between
things, however different they may appear to be, — as the wing of
a bird and the foreleg of a horse, the lungs of a bird and the swim-
bladder of a fish. The former commonly rests upon mere func-
tional, i.e. physiological, modifications ; the latter is grounded upon
structural, i.e. morphological, identity or unity. Analogy is the
correlative of physiology, homology of morphology; but the two
may he coincident, as when structures identical in morphology are
used for the same purposes and are therefore physiologically identi-
cal. Physiological diversity of structure is incessant, and continually
interferes with morphological identity of structure, to obscure or
obliterate the indications of affinity the latter would otherwise
express clearly. It is obvious that birds might he classified physio-
logically, according to their adaptive modifications or analogical
resemblances, just as readily as upon any other basis : for example,
into those that perch, those that walk, those that swim, etc. ; and,
in fact, most early classifications largely rested upon such considera-
tions. It is also evident, that when functional modifications happen
to be coincident with structural affinities, — as when the turning of
the lower larynx into a music-box coincides with a certain type of
structure, — such modifications are of the greatest service in classi-
fication, as corroborative evidence. But since all sound taxonomy
rests on morphology, on real structural affinity, we must he on our
guard against those physiological “appearances” which are pro-
verbially “deceptive.” I trust I make the principle clear to the
student. Its practical application is another matter, only to be
learned in the school of experience. This matter of

Homology op Analogy may he thus summed : Birds are
homologically related, or naturally allied or affined, according to the
sum of like structural characters employed for similar purposes ;
they are analogically related, only according to the sum of unlike
characters employed for similar purposes. A loon and a cormorant,
for instance, are closely affined, because they are both fitted in the
same way for the pursuit of their prey by flying under water. A

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PART II

dipper (family Cindidm) and a loon (family Colymlidcd) are analogous,
in so far as both are fitted to pursue their prey by flying under
water ; but they stand near opposite extremes of the ornithological
system ; they have little afiinity beyond their common birdhood —
very different structure being modified to attain the same end. So
again, conversely, the crow has vocal organs almost identical in
structure with those of the nightingale, and the organisation of the
two birds is in other respects very similar ; their afiinity or
homology is therefore close, though the crow is a hoarse croaker, the
nightingale an impassioned musician.

The Reason why Morphological Classification is so important
as to justify or even require its adoption has been very clearly
stated by Huxley, whose words I cannot do better than quote
in this connection. Speaking of animals, not as .physiological
apparatuses merely ; not as related to other forms of life and to
climatal conditions ; not as successive tenants of the earth ; but as
fabrics, each of which is built upon a certain plan, he continues :
“ It is possible and conceivable that every animal should have been
constructed upon a plan of its own, having no resemblance whatever
to the plan of any other animal. For any reason we can discover
to the contrary, that combination of natural forces which we term
Life might have resulted from, or been manifested by, a series of
infinitely diverse structures ; nor would anything in the nature of
the case lead us to suspect a community of organisation between
animals so different in habit and in appearance as a porpoise and a
gazelle, an eagle and a crocodile, or a butterfly and a lobster. Had
animals been thus independently organised, each working out its
life by a mechanism peculiar to itself, such a classification as that
now under contemplation would be obviously impossible ; a morpho-
logical or structural classification plainly implying morphological or
structural resemblances in the things classified.

“ As a matter of fact, however, no such mutual independence of
animal forms exists in nature. On the contrary, the members of the
animal kingdom, from the highest to the lowest, are marvellously
connected. Every animal has something in common with all its
fellows ; much, with many of them ; more, with a few ; and usually,
so much with several, that it differs but little from them.

“Now, a morphological classification is a statement of these
gradations of likeness which are observable in animal structures, and
its objects and uses are manifold. In the first place, it strives to
throw our knowledge of the facts which underlie, and are the cause
of, the similarities discerned, into the fewest possible general
propositions, subordinated to one another, according to their
: reater or less degree of generality ; and in this way it answers the
purpose of a memoria technica, without which the mind would be

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 105

incompetent to grasp and retain the multifarious details of anatomical
science.

“But there is a second and even more important aspect of
morphological classification. Every group in that classification is
such in virtue of certain structural characters, which are not only
common to the members of the group, but distinguish it from all
others ; and the statement of these constitutes the definition of the
group.

“Thus, among animals with vertebrae, the class Mammalia is
definable as those which have two occipital condyles, with a well-
ossified basi - occipital ; which have each ramus of the mandible
composed of a single piece of bone and articulated with the
squamosal element of the skull ; and which possess mammae and
non-nucleated red blood-corpuscles.

“But this statement of the characters of the class Mammalia
is something more than an arbitrary definition. It does not merely
mean that naturalists agree to call such and such animals Mammalia;
hut it expresses, firstly, a generalisation based upon, and constantly
verified by, very wide experience ; and, secondly, a belief arising
out of that generalisation. The generalisation is that, in nature,
the structures mentioned are always found associated together ; the
belief is that they always have been, and always will be, found
so associated. In other words, the definition of the class Mammalia
is a statement of a law of correlation, or coexistence, of animal
structures, from which the most important conclusions are deducible ”
(Introd. to Classif. of Animals^ 8vo, London, 1869, pp. 2, 3).

But broad as such laws of correlation of structure are, and
important as are the conclusions deducible, we must constantly be
on our guard against presuming upon the infallibility either of the
data or of the deduction, as the author just quoted goes on to show.
Such caution is specially required where there is no obvious reason
for the particular combination that may be found to exist. In the
case of the ostrich-like birds {Eatito}), for example, we can understand
how a flat, unkeeled breast-bone, a particular arrangement of the
shoulder -bones, and a rudimentary state of the wing -bones, are
found in combination, because all these modifications of structure
are evidently related to loss of the power of flight ; and, in point
of fact, no exception is known to the generalisation, that such
conditions of the sternal, coraco-scapular, and humeral bones always
coexist. But in all known struthious (ratite) birds, this state of
the bones in mention coexists also with a peculiar modification of
the bones of the palate, and no necessary connection between these
two sets of diverse characters is conceivable. Now, if we only
knew struthious birds, and found the combination in mention to
hold with them all, we should doubtless declare our belief that any

io6

GENERAL ORNITHOLOGY

PART II

bird having such palatal characters would also be found to possess
such imperfect wing-apparatus. But this would be going too far :
in fact, we know that the tinamous (Dromceognathce) have such a
palate, yet have a keeled sternum and functionally developed wings.
The real use and proper application of such generalisations is to
teach the lesson, that creatures exhibiting such modified combina-
tions of characters are genetically related to each other just in the
degree to which they possess characters in common, and are
genetically remote from each other in the degree to which they do
not possess characters in common : ie. that their similarities and
distinctions of structure are sure indexes of their natural affinities.
To take another case, derived from consideration of a large number
of existing birds : it is an observed fact, that a particular arrange-
ment of the plates upon the back of the tarsus, a peculiar modifica-
tion of the lower larynx or voice organ, and an undeveloped or
abortive condition of the first large feather on the hand, are found
associated in a vast series' of birds, constituting the group of Passeres
called Oscines. What possible connection there can be between
these three separate and apparently independent modifications we
cannot even surmise ; but that they have some natural and necessary
connection we cannot doubt, and that the connection is causal, not
fortuitous, is a logical inference from the observed fact, that birds
which present this particular combination are also closely related in
other structural characters ; that is, that they have all been
subjected to operative influences which have conspired to produce
the modifications observed. Given, then, a bird, with a known
oscine larynx, but unknown as to its feet and wings, it would be
a reasonable inference that these members, when discovered, would
present the characters observed to occur in like cases. But the first
lark (Alaudidce) examined would show the inference to be fallible ;
for the tarsus of such a bird is differently disposed, though a lark
has an elaborate singing apparatus, and only nine instead of ten
developed primaries. Once more : the development of a keeled
sternum, a peculiar saddle-shape of certain vertebrae, and lack of
true teeth, are characters coexisting in all the higher birds ; and,
as far as these birds are concerned, we have no hint that such a
combination is ever broken. In fact, however, the singular Creta-
ceous Ichthyornis shows us a pattern of bird in which a well-keeled
sternum and perfectly formed wings coexist with teeth in reptile-
like jaws and with fish-like biconcave vertebrae. What we learn
from this case indeed breaks down one of the most precise defini-
tions we might have made (and indeed did make) respecting birds
at large ; but in its failure we are taught how great is the modifica-
tion of geologically recent birds from their primitive generalised
ancestry ; we learn something likewise of the steps of such

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 107

modification, and of the length of time required for the process.
It is the history of attempts to frame definitions of groups in
zoology, that they are all liable to be negatived by new discoveries,
and therefore to be broken down and require remodelling as our
knowledge increases. It is to he readily perceived that the
ability to draw distinctions and make definitions of groups is as
much the gauge of our ignorance as the test of our knowledge ; for
all groups, like all species, come to he such by modification so
gradual, so slight in each successive increment of difference, that, if
all the steps of the process were before our eyes, we should he able
to limit no groups whatever in a positive, unqualified manner. All
would merge insensibly into one another, he inseparably linked in
as many series as there have been actual lines of evolutionary
progress, and finally converge to the one or few starting-points of
organised beings.

Practically, however, the case is quite the reverse, — happily for
the comfort of the working naturalist, however sadly the philosopher
may deplore the ignorance implied. Degrees of likeness and
unlikeness do exist, which when rightly interpreted enable us to
mark off groups of all grades with much facility and precision, and
thus erect a morphological classification which recognises and defines
such degrees, and explains them upon the principles of Evolution.
The way in which the principles of such classification are to he
practically applied gives occasion for some further remarks upon

Zoolog’ieal Chapacters. — A character,” in zoological language,
is any point of structure which may he perceived and described for
the purpose of comparing or contrasting animals with one another.
Thus, the conditions of the sternum, palate, tarsus, larynx, as
noted in preceding paragraphs, are each of them characters
which ma}^ be used in describing individual birds, or in framing
definitions of groups of birds. Morphological characters, with
which the classification we have adopted alone concerns itself, may
be derived from the structure of a bird considered in any of its
relations, or as affected by any of the conditions to which it is
subjected. Thus emlryological characters are those afforded by the
bird during the progress of its development in the egg,, from the
almost structureless germ to the fully formed chick. Such
characters of the embryo in its successive stages are of the utmost
significance ; for it is a fact that the germ of each of the higher
organisms goes through a series of developmental changes which,
at each succeeding step in the unfolding of its appropriate plan of
structure, causes it to resemble the adult state of animals lower
than itself in the scale of organisation. In fine, the history of the
evolution of every individual bird epitomises the history of those
changes which birds collectively have undergone in becoming what

io8

GENERAL ORNITHOLOGY

PART II

they are by modified descent from lower organisms. Such transitory
stages of any embryo, therefore, give us glimpses of those evolu-
tionary processes which have affected the group to which it belongs.
Any bird, for example, when a germ, is at first on the plane of
organisation of the very lowest known creatures, — one of the
Protozoa^ or single-celled animals. As its germ develops, and its
structure becomes more complicated by the formation of parts and
organs successively differentiated and specialised, it rises higher
and higher in the scale of being. At a certain stage very early
reached (for the steps by which it becomes like any invertebrate are
very speedily passed over) it resembles a fish in possessing gill-like
slits, several aortic arches, no true kidneys, no amnion, etc. Further
advanced, losing its gills, gaining kidneys and amnion, etc., it rises
to the dignity of a reptile, and at this stage it is more like a
reptile than like a bird ; having, for example, a number of separate
bones of the wrist and ankle, no feathers, etc. The assumption of
its own appropriate characters, i.e. those by which it passes from a
reptilian creature to become a bird, is always the last stage reached.
We can thus actually see and note, inside any egg-shell, exactly
those progressive steps of development of the individual bird which
we believe to have been taken on a grand scale in nature for the
evolution of the class Aves from lower forms of life ; and the lesson
learned is fraught with significance. It is nothing less than the
demonstration in ontogeny (genesis of the individual) of that
jphylogeny (genesis of the race) by which groups of creatures come
to be. The interior of any adult bird, again, furnishes us with all
kinds of ordinary anatomical characters, derived from the way we
perceive the different organs and systems of organs to be fashioned
in themselves, and arranged with reference to one another. The
finishing of the outward parts of a bird gives us the ordinary
external characters, in the way in which the skin and its appendages
are modified to form the covering of the bill and feet, and to fashion
all kinds of feathers. Birds being of opposite sexes, and such
difference being not only indicated in the essential sexual organs,
but usually also in modifications in size or shape of the body or
quality of the plumage and other outgrowths, a set of sexual
characters are at our service. Birds are also sensibly modified in
their outward details of feathering by times of the year when the
plumage is changed, and this renders appreciation of seasonal
characters possible. All such circumstances, and others that could
be mentioned, such as effects of climate, of domestication, etc., in so
far as they in any way affect the structure of birds, conspire to
produce zoological characters, as these are above defined. Such
characters, according as they result from more or less profound
impressions made upon the organism, are of more or less “ value ”

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 109

in taxonomy ; being of all grades, from the trivial ones that serve
to distinguish the nearest related species or varieties, to the
fundamental ones that serve to mark off primary divisions. Thus
the “ character ” of possessing a backbone is common to all animals
of an immense series called Vertebrata. The “ character ” of feathers
is common to all the class Aves ; of toothless jaws to all modern
birds ; of a keeled sternum to all the sub-class Carinatce ; of feet
fitted for perching to all the order Passeres ; of a musical apparatus
to all the sub-order 0 seines ; of nine primaries to all the family
FringilUdce ; of crossed mandibles to all of the genus Loxia ; of
white bands on the wings to all of the species Loxia leucoptera.
There is thus seen a sliding scale of valuation of characters, from
those involving the most profound or primitive modifications of
structure to those resting upon the most superficial or ultimate
impressions. It will also be obvious that every ulterior modifica-
tion presupposes inclusion of all the prior ones ; for a white-
winged crossbill, to be itself, must be a loxian, fringilline, oscine,
passerine, carinate, modern, avian, vertebrated animal. The more
characters, of all grades, that any birds share in common, the more
closely are they related, and conversely. Obviously, the possession
of more or fewer characters in common results in

Degrees of Likeness. — Were all birds alike, or did they all
differ by the same characters to the same degree, no classification
would be possible. It is a matter of fact that they do exhibit all
degrees of likeness possible within the limits of their Avian nature ;
it is a matter of belief that these degrees are the necessary result of
Evolution — of descent with modification from a common ancestry ;
and that, being dependent upon that process, they are capable of
explaining it if rightly interpreted. For example : Two white-
winged crossbills, hatched in the same nest, scarcely differ percep-
tibly (except in sexual characters) from each other, and from the
pair that laid the eggs. We call them “ specifically ” identical ;
and the sum of the differences by which they are distinguished
from any other kinds of crossbills is their “ specific character.” All
the individual crossbills which exhibit this particular sum consti-
tute a “ species.” In this case, the genetic relationship of offspring
and parent is unquestionable — ^it is an observed fact. Now turn
to the extremely opposite case. The difference between our cross-
bills and the Cretaceous Ichthyornis is enormous : I suppose it is
nearly the greatest known to subsist between any two birds what-
soever. But the Ichthyornis and the Loxia are also separated by
a corresponding!}^ immense interval of time^ and presumably by
correspondingly enormous differences in conditions of environment —
in their physical surroundings. It is a logical inference that these
two things — difference in physical structure and difference in

no

GENERAL ORNITHOLOGY

PART II

physical environment — are in some way correlated and co-ordinated.
If we presume, upon the theory of evolution, that, despite the great
difference, a crossbill is genetically related to some such -bird as an
Ichthyornis, as truly as it is to its actual parents, only much more
remotely, and that the difference is due to modifications impressed
upon its stock in the course of time, conformably with changing
conditions of environment, we shall have a better explanation of
the difference than any other as yet offered — an explanation, more-
over, which is corroborated by all the related facts we know, and
with which no known facts are irreconcilable. But to correctly
gauge and formulate the degrees of likeness or unlikeness between
any two birds is to correctly “ classify ” them ; and if these degrees
rest, as we believe they do, upon nearness or remoteness of genetic
relationship, classification upon such basis becomes the truest
attainable formulation of “ natural affinities.” It is the province of
morphological classification to search out those natural affinities
which the structure of birds indicates, and express them by divid-
ing birds into groups, and subdividing these into other groups, of
greater or lesser value or grade, according to the fewer or more
characters shared in common, — that is, according to degrees of like-
ness ; that is, again, according to genealogical relationship or con-
sanguinity.

Zoological Groups. — To carry any scheme of classification into
practical effect, naturalists have found it necessary to invent and apply
a system of grouping objects whereby the like may come together and
be separated from the unlike. They have also found it expedient to
give names to all these groups, of whatever grade, such as class,
order, family, genus, species, etc. ; and to stamp each such group with
the value of its grade, or its relative rank in the scale, so that
it may become currency among naturalists. The student must

observe, in the first place, that the value of each such coinage is
wholly arbitrary, until sanctioned and fixed by common consent.
The term “ class,” for example, simply indicates that naturalists
agree to use that word to designate a conventional group of a
particular grade or value. Indispensable as is some such acceptable
medium of exchange of ideas among naturalists, their groups are
not fixed, have no natural value, and in fact have no actual exist-
ence in the treasury of Nature. It cannot be too strongly impressed
upon the student that Nature makes no bounds, — Natura non facit
saltus ; there are no such abrupt transitions in the unfolding of
Nature’s plan, no such breaks in the chain of being, as he would be
led to suppose by our method of defining and naming groups. He
must consider the words “ class,” “ order,” etc., as wholly arbitrary
terms, invented and designed to express our ideas of the relations
which subsist between any animals or sets of animals. Thus, for

SEC. II PRINCIPLES AND PRACTICE OP CLASSIFICATION iii

example, by the term the “ Class of Birds ” we signify simply the
kind and degree of likeness which all birds share, such being also
the kind and degree of their unlikeness from any other animals ;
the word “ class ” being simply the name or handle of the general-
isation we make respecting their relations with one another and
with other animals ; it represents an abstract idea, is the expression
of a relation. True, all birds embody the idea ; but “ class ” is
nevertheless an abstraction. Now, as intimated earlier in this
essay, the definition of the idea we attach to the term — the limita-
tion of the class Aves — depends entirely upon how much we know
of the relation intended to be expressed. It so happens that no
animals are known which cannot be decided to belong, or not to
belong, to the conventional class of birds, because we have found it
convenient and expedient to consider the presence of feathers a fair
criterion or necessary qualification. But what, when an animal is
discovered the covering of whose body is half-way between the
scales of a lizard and the plumes of a bird, and whose structure is
otherwise as equivocal This may happen any day. A feather is
certainly a modified scale ; a feather has doubtless been developed
out of a scale. In the case supposed, we should have to modify our
definition of the “ Class of Birds ” ; that is, change our ideas upon
the subject, and alter the boundary-line we established between the
classes of birds and reptiles ; whereas, were a “ class ” something
naturally definite, independent, and fixed, all that we could learn
about it would only tend to establish it more surely. The same
obscurity and uncertainty of definition attaches to groups of every
grade — from the Animal “ Kingdom ” itself, which cannot be cut
clear of the Vegetable “Kingdom” — down through classes, orders,
families, genera, species, and varieties — yes, to the individual itself,
which, however unmistakable among higher organisms, cannot always
be predicated of the lowermost forms of Life. Such divisions, of
whatever grade, as we are able to establish for the purposes of
classification, depend entirely upon the breaks and defects in our
knowledge. There is no such thing as drawing hard-and-fast
lines anywhere, for none such exist in Nature.

Taxonomic Equivalence of Groups. — But, however arbitrary
they may be, or however obscure or fluctuating may be their
boundaries, groups we must have in zoology, and groups of different
grades, to express different degrees of likeness of the objects
examined, and so to classify them. It is a great convenience,
moreover, to have a recognised sliding-scale of valuation of groups
from the highest to the lowest, and an accepted valuation. Just as
in a thermometric scale, there are degrees designated as those of
the boiling-point of water, the heat of the blood, the freezing of
water, of mercury, etc.; so there are certain degrees of likeness

II2

GENERAL ORNITHOLOGY

PART II

conventionally designated as those of dass^ order, family, genus, and
species; always accepted in the order here given, from higher to
lower groups. (There are various others, and especially a number
of intermediate groups, generally distinguished by the prefix sul-, as
sub-family ; but those here given are generally adopted by English-
speaking naturalists, and suffice to illustrate the point I wish to
make.) It may sound like a truism to say that groups of the same
grade, bearing the same name, whatever that may be, must be of
the same value, — must be based upon and distinguished by charac-
ters of equal or equivalent importance. Equivalence of groups is
necessary to the stability and harmony of any classificatory system.
It will not do to frame an order upon one set of characters here,
and there a family upon a similar set of characters ; but order must
differ from order, and family from family, by an equal or corre-
sponding amount of difference. Let a group called a family differ as
much from the other families in its own order as it does from some
other order, and by this very circumstance it is not a family but an
order itself. It seems a very simple proposition, but it is too often
ignored, and always with practical ill result. Two points should be
remembered here : First, that absolute size or numerical bulk of a
group has nothing to do with its taxonomic value : one order may
contain a thousand species, and another be represented by a single
species, without having its ordinal valuation affected thereby.
Secondly, any given character may assume different importance, or
be of different value, in its application to different groups. Thus,
the number of primaries, whether nine or ten, is a family charac-
ter almost throughout Oscines; but in one oscine family (Fireonidce)
it has scarcely generic value. It is difficult, however, to determine
such a point as this without long experience. Nor is it possible, in
fact, to make our groups correspond in value with entire exactitude.
The most we can hope for is a reasonable approximation. As in the
thermometric simile above given, “blood-heat” and other points
fluctuate, so does order not always correspond with order, nor
family with family, in actual significance. What degree of differ-
ence shall be “ordinal” ? What shall be a difference of “family”?
What shall be “generic” and what “specific” differences? Such
questions are more easily asked than answered. They demand
critical consideration.

Valuation of Charaeters. — In a general way, of course, the
greater the difference between any two objects, the more “ import-
ant” or “fundamental” are the “characters” by which they are
distinguished. But what makes a character “important” or the
reverse ? Obviously, what it signifies represents its importance.
We are classifying morphologically, and upon the theory of Evolu-
tion ; and in such a system a character is important or the reverse.

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 113

simply as an exponent of the principles, or an illustration of the
facts, of evolutionary processes of Nature, according to the unfold-
ing of whose plans of animal fabrics the whole structure of living
beings has been built up. Why is the possession of a backbone
such a “fundamental” character that it is used to establish one of
the primary branches of the animal kingdom '? It is not because so
many millions of creatures possess it, but because it was introduced
so early in the evolutionary process, and because its introduction led
to the most profound modification of the whole structure of the
animals which became possessed of a vertebral column. Why is
the possession by a bird of biconcave vertebrae so significant ? Not
because all modern birds have saddle-shaped vertebrae, but because
to have biconcave vertebrae is to be quoad hoc fish-like. Why is pre-
sence or absence of teeth so important? Not that teeth served
those old birds better than a horny beak serves modern ones, but
because teeth are a reptilian character. Obviously, to be fish-like
or reptile-like is to be by so much unbirdlike ; the degree of differ-
ence thus indicated is enormous ; and a character that indicates such
degree of difference is proportionally “important” or “fundamental,”
— just what we were after. By knowledge of facts like these, and
by the same process of reasoning, a naturalist of tact, sagacity, and
experience is able to put a pretty fair valuation upon any given
character ; he acquires the faculty of perceiving its significance, and
according to what it signifies does it possess for him its taxonomic
importance. As a matter of fact, it seems that characters of all
sorts are to be estimated chronologically. For, if animals have come
to be what they are by any process that took time to be accom-
plished, the characters earliest established are likely to be the most
fundamental ones, upon the introduction of which the most import-
ant train of consequences ensue. Feathers, for example, as the
Archmopteryx teaches us, were in full bloom in the Jurassic period,
and they are still the most characteristic possession of birds : all
birds have them ; they are a class character. If they had been
taken on quite recently, we may infer that many creatures otherwise
entirely avian might not possess them, and they would have in
classification less significance than that now rightly attributed
to them. On the other hand, we cannot suppose that the finishing
touches, by which, in the presence of white bands on the wings of
Loxia leucoptera, and their absence in Loxia curvirostra, these two
“ species ” are distinguished, were not very lately given to these
birds. It is a very late step in the process, and correspondingly in-
significant ; it is of that value or importance which we call “specific.”
The same method of reasoning is available for determining the
value of any character whatever, and so of estimating the grade of
the group which we establish upon such character. As a rule,
I

GENERAL ORNITHOLOGY

PART II

II4

therefore, the length of time a character has been in existence, and
its taxonomic value, are correlated, and each is the exponent of the
other.

“ Types of Stpueture.” — In no department of natural history
has the late revolution in biological thought been more effective
than in remodelling, presumably for the better, the ideas underlying
classification. In earlier days, when “ species ” were supposed to be
independent creations, it was natural and almost inevitable to
regard them as fixed facts in nature. A species was as actual and
tangible as an individual, and the notion was, that, given any two
specimens, it should be perfectly possible to decide whether they
were of the same or different species, according to whether or not
they answered the “ specific characters ” laid down for them. The
same fancy vitiated all ideas upon the subject of genera, families,
and higher groups. A “ genus ” was to be discovered in nature,
just like a species ; to be named and defined. Then species that
answered the definition were “ typical ” ; those that did not do so
well were “ sub-typical ” ; those that did worse were “ aberrant.” A
good deal was said of “types of structure,” much as if living crea-
tures were originally run into moulds, like casting type-metal, to
receive some indelible stamp ; while — to carry out my simile — it
was supposed that by looking at some particular aspect of such an
animal, as at the face of a printer’s type, it could be determined in
what box in the case the creature should be put ; the boxes them-
selves being supposed to be arranged by Nature in some particular
way to make them fit perfectly alongside each other by threes or
fives, or in stars and circles, or what not. How much ingenuity
was wasted in striving to put together such a Chinese puzzle as
these fancies made of Nature’s processes and results, I need not
say ; suffice it, that such views have become extinct, by the method
of natural selection, and others, apparently better fitted to survive,
are now in the struggle for existence. Eightly appreciated, how-
ever, the expression which heads this paragraph is a proper one.
There are numberless “ types of structure.” It is perfectly proper
to speak of the “ vertebrate type,” meaning thereby the whole plan
of organisation of any vertebrate, if we clearly understand that such
a type is not an independent or original model conformably with
which all backboned animals were separately created, but that it is
one modification of some more general plan of organisation, the un-
folding of which may or did result in other besides vertebrated
animals ; and that the successive modifications of the vertebrate
plan resulted in other forms, equally to be regarded as “types,” as
the reptilian, the avian, the mammalian. Upon this understanding,
a group of any grade in the animal kingdom is a “ type of struc-
ture,” of more general or more special significance, presumably

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 115

according to the longer or shorter time it has been in existence.
An individual specimen is “ typical ” of a species, a species is
“typical” of a genus, etc., if it has not had time enough to be
modified away from the characters which such species or genus
expresses. Any set of individuals, that is, any progeny, which
become modified to a degree from their progenitors, introduce a new
type ; and continually increasing modification makes such a type
specific, generic, and so on, in succession of time. There must have
been a time, for example, when the Avian and Reptilian “ types ”
began to diverge from each other, or, rather, to branch apart from
their common ancestry. In the initial step of their divergence,
when their respective types were beginning to be formed, the differ-
ence may have been infinitesimal. A little farther along, the incre-
ment of difference became, let us say, equivalent to that which serves
to distinguish two species. Wider and wider divergence increased
the difference, till genera, families, orders, and finally the classes
of Eeptilia and Aves, became established. In one sense, therefore, —
and it is the usual sense of the term, — the “type” of a bird is that
one which is farthest removed from the reptilian type, — which is most
highly specialised by differentiation to the last degree from the char-
acters of its primitive ancestors. One of the Oscines, as a thrush or
sparrow, would answer to such a type, having lost the low, primi-
tive, generalised structure of its early progenitors, and acquired very
special characters of its own, representing the extreme modification
which the stock whence it sprang has undergone. In a broader
sense, however, the type of a bird is simply the stock from which it
originated : and in such sense the highest birds are the least typical,
being the farthest removed and the most modified derivatives of
such stock, the characters of which are consequently remodelled and
obscured to the last degree. Two opposite ideas have evidently
been confused in the use of the word “ Type.” They may be dis-
tinguished by inventing the word teleotype (Gr. reAeo?, teleos, final,
i.e. accomplished or determined ; formed like teleology, etc.) in the
usual sense of the word type ; and using the word we already
possess, prototy;pe (Gr. irpOtros, protos, first, leading, determining), in
the broader sense of the earlier plan whence any teleotype has been
derived by modification. Thus, Ichthyornis or Archceopteryx is proto-
typic of modern birds, any of which are teleotypic of their ancestors.
It may be further observed that any form which is teleotypic in its
own group is prototypic of those derived from it. Thus, the
Archceopteryx, so prototypic of modern birds, was a very highly
specialised teleotype of its own ancestry. A little reflection will
also make it clear that the same principle of antitypes (opposed
types) is applicable to any of our groups in zoology. Any group is
teleotypic of the next greater group of which it is a member ; prototypic of

ii6

GENERAL ORNITHOLOGY

PART II

the next lesser one. Any species is teleotypic of its genus ; any genus,
of its family ; any family, of its order ; and conversely ; that is to
say, any species represents one of the ulterior modifications of the
plan of its genus. The Class of Birds, for example, is one of the
several teleotypes of Vertebrata, i.e. of the vertebrate plan of struc-
ture ; representing, as it does, one of several ways in which the
vertebrate prototype is accomplished. Conversely, the Class of
Birds is prototypical of its several orders, representing the plan
which these orders severally unfold in different ways. And so on,
throughout any series of animals, backward and forward in the
process of their evolution : any given form being teleotypic of its
predecessors, prototypic of its successors. All existing forms are
necessarily teleotypic, — only prototypic for the future. Prototype,
in the sense here conveyed, indicates what is often expressed by the
word archetype. But the latter, as I understand its use by Owen
and others, signifies an ideal plan never actually realised ; the
“ archetype of the vertebrate skeleton,” for example, being some-
thing no vertebrate ever possessed, hut a theoretical model — a
generalisation from all known skeletons. The correspondence of my
use of “prototypic” with a common employment of “ archetypic,”
and of “teleotypic” as including both “attypic” and “ etypic,” is
noted below. ^

The actual and visible genetic relationships of living forms being
practically restricted to individuals of the same species, — parents
and offspring specifically identical, — it would seem at first sight
that species must be the modified descendants of their respective
genera, in order to he teleotypic of any such next higher group.
But nothing descends from a genus, or any other group ; every-
thing descends from individuals ; a genus, like any other group,
is an abstract statement of a relation, not a begetter of anything.
To illustrate : the “ genus Turdus ” is represented, let us say, by a
score of species : if these species he rightly allocated in the genus,
they are all the modified descendants of a form which was, before
they severally branched off, a specific form; and the “genus

^ ‘^Archetypical characters are those which a group derives from its progenitor,
and with which it commences, but which in much modified descendants are lost ;
such, for example, is the dental formula of the Educabilia (M | PM f C ^ 1 f x 2),
— a formula, as shown by Owen, very prevalent among early members of the group,
but generally departed from more or less in those of the existing faunas. Attypical
characters are those to the acquisition of which, as a matter of fact, we find that
forms, in their journey to a specialised condition, tend. . . . Etypical characters are
exceptional ones, and which are exhibited by an eccentric offshoot from the common
stock of a group ” (Gill, Pr. Am. Assoc. Adv. Sci., xx. 1873, p. 293). To illustrate
in birds : A generalised lizard-like type of sternum is archetypic of any bird’s ster-
num. The sternum of the lizard-like animals whence birds actually descended is
prototypic ; the keeled sternum of a carinate bird is attypical in most birds, etypical
in the peculiar state in which it is found in Stringops ; but equally teleotypic in both
instances.

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 117

Turdus ” in the abstract is simply that form ; and that form is
prototypic of its derivatives. In the concrete, as represented
by its teleotypes, the genus Turdus sums the modifications which
these have collectively undergone, without specifying the particular
modifications of any of them ; it expresses the way in which they
are all like one another, and in which they are all unlike the re-
presentatives of any other genus. Thus what is above advanced
is seen to hold, though genera and all other groups are actual
descendants of individuals specifically identical.

Generalised and Specialised Forms. — Taking any one group
of animals — say the genus Turdus, of numerous species — and con-
sidering it apart from any other group, we perceive that it represents
a certain assemblage of characters peculiar to itself, aside from those
more fundamental ones it includes of its family, order, etc. Its
particular characters we call “generic.” Among the numerous
teleotypic forms it includes, there is a wide range of specific varia-
tion, within the limits of generic relationship. Some of its species
are modified farther away than some others are from the generic
standard or type to which all conform more or less perfectly. The
former, having more peculiarities of their own, are said to be the
most specialised ; the latter, having fewer peculiarities, are the least
specialised. Those that are the least specialised are obviously the
most generalised ; and this means that we believe them to be nearest
to the stock whence all have together descended with modification.
The application of this illustration to great groups shows us the
principle upon which any form is said to be generalised or specialised.
The IchtJiyornis, with its fish-like vertebrae, reptile-like teeth, bird-
like sternum and shoulder-girdle, is a very generalised form. A
thrush is the opposite extreme of a highly specialised form. The
two are also separated by an enormous interval of time ; one being
very old, the other quite new ; a chronological sequence is here
perceived. Since the evolutionary processes concerned in the modi-
fication on the whole represent progress from simplicity to com-
plexity of organisation, and therefore ascent in the scale of
organisation, a generalised type, an ancient type, and a simple type
are on the whole synonymous, and to be contrasted with forms
specialised, recent, and complex. They therefore respectively corre-
spond to

“ Low ” and “ High ” in the Seale of Organisation. — All

existing birds are very closely related, notwithstanding the great
numerical preponderance of the class in the present geological
epoch. This outbreak, as it were, of birds upon the modern scene,
is like the nearly simultaneous bursting into bloom of a mass of
flowers at the end of one branch of the Sauropsidan stem. All
modern birds, inTact, are strongly specialised forms, so much so that

ii8

GENERAL ORNITHOLOGY

PART II

it is difficult to predicate “ high ” or “ low” within such a narrow
scale. The great group Passeres^ for example, comprehending a
majority of all known birds, is scarcely more different from other
birds than are the families of reptiles from each other, and among
Passeres we have little to go upon in deciding “ high ” or “ low ”
beyond the musical ability of Oscines. It is hard to see much
difference in actual complexity of organisation between those birds
regarded as the lowest, as an ostrich or a penguin, and those con-
ceded to be highest, as a swallow or sparrow. Nevertheless, in a
larger perspective, as between a fish, a reptile, and a bird, the
student will readily perceive the hearing of the ideas attached to
the terms “ low ” and ‘‘ high ” in the scale of organisation. Creatures
rise in the scale by a number of correlated modifications and in the
course of time (for it takes time to evolve a class of birds from
sauropsidan stock as really as it does to develop the germ of an egg
into the body of a chick). Progressive differentiation and specialisa-
tion of structure and function in due course elaborates diversity
from sameness, complexity from simplicity, the “ high ” special from
the “ low ” general plan of organisation ; the culmination in man of
the vertebrate type, first faintly foreshadowed in the embryonic
Ascidian. No one should venture to foretell the result of infinit-
esimal increments in elevation of structure and function, nor pre-
sume to limit the infinite possibilities of evolutionary processes,
either in this actual world or in a foretold next one.

As to “ evidences of design ” in the plan of organised beings, it
may be said simply that every creature is perfectly “ designed ” or
fitted for its appropriate activities, and perfectly adapted to its
conditions of environment. In fact, it must be so fitted and
adapted, or it would perish. Whether it so determines itself, or is
so determined, is a teleological question. The truth remains that
every creature is perfect in its own way. A worm is as perfectly
fitted to be a worm, as is a bird to be a bird ; in fact, were it not,
it would either turn into something else, or cease to be. A spade
is as perfect an organisation of the spade kind, as is the steam-engine
of that kind of an organisation ; though the difference in complexity
of structure and functional capacity, like that between the lowly
organised ascidian generality and the highly organised avian
speciality, is enormous.

One word more : The class of mammals is highest in the scale
of organisation. The class of birds is next highest. But it does
not follow, from this relation sustained by Mammalia and Aves
collectively, that every mammal must be more highly organised
than every bird. It is difficult to say how a mole or a mouse is a
more elaborate or more capable creature than a canary-bird, physic-
ally or mentally. The relative rank of two groups is determined

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 119

by balancing the aggregate of their structural characters. In large
series, the average of development, not the extremes either way, is
taken into account ; so that the lowest members of a higher group
may be below the highest members of the next lower group. The
common phrase, “ below par,” or “ above par,” is most applicable
to such cases.

Maehinepy of Classification. — The inexperienced student may
be glad to be given some explanation of the way in which the
taxonomic principles we have discussed are applied, and carried into
practical effect in classifying birds. Our machinery for that purpose
is our inheritance from those naturalists who held very different
views from those which touch the evolutionary key-note of modern
classification. It is clumsy, and does not work well as a means of
expressing the relations we now believe to be sustained by all organ-
isms toward one another ; but it is the best we have. Systematic
zoology, or the practice of classification, has failed to keep pace with
the principles of the science ; we are greatly in need of some new
and sharper “ tools of thought,” which shall do for zoology what the
system of symbols and formulae has done for chemistry. fTe want
some symbolic formulation of our knowledge. The invention of a prac-
ticable scheme of classification and nomenclature, which should
enable us to formulate what we mean by Turdus viscivorus, as a
chemist symbolises by SO^H2 what he understands hydrated sul-
phuric acid to be, would be an inestimable boon to working
naturalists. The mapping out of groups with connecting lines to
indicate their genetic relations, in the form of the phylum, is a
common practice ; but that, like any other pictorial representation
of a “ family tree,” is not the graphic symbolisation required. The
first steps in this direction have been tentatively taken already by
the late Mr. A. H. Garrod and others : we already have a mother
of the required invention in the necessity of the case, and may hope
that the father will not be long in coming.

Under the present system. Birds are called a “Class” of Verte-
brates, and are subdivided into “orders,” “families,” “genera,”
“ species,” and “ varieties,” as already sufficiently indicated. Groups
intermediate to any of these may be recognised ; and if so, are
usually distinguished by the prefix sub-. Many other terms are
in occasional use, as “ tribe,” “ race,” “ series,” “ cohort,” “ super-
family ” ; but the six first mentioned are the best established ones
among English-speaking naturalists. Their sequence is fixed, as
above, from higher to lower, in relative rank.^ With the exceptions

^ The expression “higher group,” in the sense of relative rank in the taxonomic
scale, will of course he distinguished from the same expression when applied to the
relative rank in the scale of organisation of the objects classified. An order of birds
is a “ higher group ” than a family of birds, in the former sense, but no higher than
an order of worms, in the latter sense.

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GENERAL ORNITHOLOGY

PART II

to be presently noted, the names of groups are arbitrary, at the will
of the person who establishes and designates them. The framer of
a genus, or the describer of a species, calls it what he pleases, and
the name he gives holds, subject to certain statutory regulations
which naturalists generally agree to abide by. The exceptions are
the names of families and sub-families, the former commonly being
made to end in -idm^ the latter in -ince : family Turdidce ; sub-family
Turdince. This is a great convenience, since we always know the
rank intended to be noted by these word -forms. The names of
groups higher than species are almost invariably single words ; as,
order Fasseres ; but sometimes, especially in cases of intermediate
groups, two words are used, one qualifying the other ; as, sub-order
Fasseres Acromyodi, or oscine Passeres. A generic or sub-generic
name is always a single word ; these, and the names of all higher
groups, invariably begin with a capital letter.

Until quite recently, the scientific name of any individual bird
almost invariably consisted of two terms, generic and specific, — the
name of the genus, followed by the name of the species j as, Turdus
viscivorus, for the missel -thrush. This is the “binomial nomen-

clature ” (badly so called, for “ binominal ” or “ bionymic ” would be
better) ; introduced by Linnaeus in the middle of the last century.
It was a great improvement upon the former method of giving either
single arbitrary names to birds, often a mere Latin translation of
their vernacular nickname, or long descriptive names of several
words ; probably no other single improvement in a method of
nomenclature ever did so much to make the technique of nomen-
clature systematic. To couple the two terms at all was a great
thing, the convenience of which we who never felt its want can
hardly appreciate. To follow the generic by the specific term was
itself of the same advantage that it is to have the Smiths and
Browns of a directory entered under S and B, instead of by Johns
and Jameses ; besides according with the genius of the Eomance
languages, which commonly put the adjective after the noun. A
Frenchman, for example, would say, Bec-crois6 aux ailes Handles de
VAmirique septentrionale, or “ Bill-crossed to the wings white of the
America north,” where we should say, “North American white-
winged Crossbill,” and Linnaeus would have written Loxia leucoptera.
The binomial scheme worked so well that it came to have the
authority and force of a statute, which few subsequent naturalists
have been inclined, and fewer have ventured, to violate ; while it
became an ex post facto law to prior naturalists, ruling them out of
court altogether, as far as the legitimacy of any of the names they
had bestowed was concerned. It necessarily rested, however, or at
any rate proceeded upon, the false idea of a species as a fixity.
Linnaeus himself experienced the inadequacy of his system to deal

SEC. II PRINCIPLES AND PRACTICE OF CLASSIFICATION 121

binomially with those lesser groups than species, commonly called
“ varieties,” now better designated as “ conspecies ” or “ subspecies ” ;
and he often used a third word, separated however from the binomial
name by intervention of the sign “var.” or some other symbol.
Thus, if he had supposed an American crossbill to be a variety of a
European Loxia leucoptera, he might have called it Loxia leuco'ptera, a,
americana. Some years ago, in treating of this subject, I urged the
necessity of recognising by name a great number of forms of our
birds intermediate between nominal species, and connecting the
latter by links so perfect, that our handling of species required
thorough reconsideration. The dilemma arose, through our very
intimate knowledge of the climatic and geographical variation of
species, either to discard a great number that had been described,
and so ignore all the ultimate modifications of our bird-forms ; or else
to recognise as good species the same large number of forms that we
knew shaded into each so completely that no specific character could
be assigned. In the original edition of the Key to North American
Birds (1872), I compromised the matter by reducing to the rank of
varieties the nominal species that were known or believed to inter-
grade ; and the original edition of my Check List (1873) distinguished
such by the sign “ var.” intervening between the specific and the
subspecific name. I subsequently determined to do away with the
superfluous term “var.,” and in the next edition of the Check List
(1882) reverted to a purely trinomial system of naming the equi-
vocal forms as, Loxia curvirostra americana. This system is found
to work well, and seems likely to come into general use.^

The Student cannot be too well assured that no such things
as species, in the old sense of the word, exist in nature, any more
than have genera or families an actual existence. Indeed they
cannot be, if there is any truth in the principles discussed in our
earlier paragraphs. Species are simply ulterior modifications, which
once were, if they be not still, inseparably linked together ; and
their nominal recognition is a pure convention, like that of a genus.
More practically hinges upon the way we regard them than turns
upon our establishment of higher groups, simply because upon the
way we decide in this case depends the scientific labelling of specimens.
If we are speaking of a robin, we do not ordinarily concern our-
selves with the family or order it belongs to, but we do require a

^ Since the above was penned, the trinominal or trionymic system of nomenclature
has been formulated and fully adopted by the committee on Nomenclature of
the American Ornithologists’ Union, of which Dr. Cones was chairman ; and the
decision of that body of nomenclatural legislators, as expressed in its Canons of
Nomenclature, has been recognised as authoritative not only by American ornitholo-
gists in general, but by naturalists in other departments of zoology, notably mam-
malogy, herpetology, ichthyology, malacology, and entomology. The scheme has
become well known to British ornithologists as a distinctive feature of the “ American
School.”

122

GENERAL ORNITHOLOGY

PART II

technical name for constant use. That name is compounded of its
genus, species, and variety. No infallible rule can be laid down for
determining what shall be held to be a species, what a conspecies,
subspecies, or variety. It is a matter of tact and experience, like
the appreciation of the value of any other group in zoology. There
is, however, a convention upon the subject, which the present
workers in ornithology in America find available ; and there is no
better rule to go by. They treat as “ specific ” any form, however
little different from the next, that is not known or believed to inter-
grade with that next one; between which and the next one no
intermediate equivocal specimens are forthcoming, and none, con-
sequently, are supposed to exist. This is to imply that the differen-
tiation is accomplished, the links are lost, and the characters actually
become “specific.” They treat as “varietal” of each other any
forms, however different in their extreme manifestation, which they
know to intergrade, having the intermediate specimens before them,
or which they believe with any good reason do intergrade. If the
links still exist, the differentiation is still incomplete, and the
characters are not specific, but only varietal, in the literal sense of
these terms. In the latter case, the oldest name is retained as the
specific one, and to it is appended the varietal designation : as,
Turdus migratorius propinquus. The specific and subspecific words
are preferably written with a small initial letter, even when derived
from the name of a person or place, after the example of Dr. P. L.
Sclater and other eminent British naturalists.

One other term than those just considered sometimes forms part
of a bird’s scientific name : this is the subgenus. When introduced,
it always follows the generic term, in parentheses ; thus, Turdus
{Merula) torquatus. This is cumbrous, especially when there are
already three terms, and is little used. I have latterly discarded it
altogether. There is no difference in kind between a subgenus and
a genus, — it is a difference of slight degree merely ; and modern
genera have so multiplied that one can easily find a single name for
any generic refinement he may wish to indulge.

It has always been customary to write after the bird’s name the
name of the original describer of the species, — originally and
properly, as the authority or voucher for the validity of the species
named. But as genera multiplied, it was often found necessary to
change the generic name, the species being placed in another genus
than that to which its original namer referred it. The name of the
person who originated the new combination came to be generally
suffixed, presumably as the authority for the validity of the classi-
fication implied. As this was to ignore the proprietorship of the
original describer, it became customary to retain that describer’s
name in parentheses and add that of the classifier ; thus, Turdus

SEC. Ill

EXTERIOR PARTS OF BIRDS

123

migmtorius Linnaeus ; Planesticus migratorius (Linn.) Bonaparte. The
practice still prevails.

It would take me too far to go fully into the rules of nomen-
clature : some few points may be noted. A proper sense of justice
to the describers of new genera, species, and varieties prompts us to
preserve inviolate the names they see fit to bestow, with certain
salutary provisions. Hence arises the “law of priority.” The first
name given since 1758 is to be retained and used, if it can be iden-
tified with reasonable certitude ; that is, if we think we know what
the giver meant by it. But it is to be discarded, and the next name
in priority of time substituted, if it is “ glaringly false or of express
absurdity,” — as calling an English bird “ africanusfi or a black one
alhusP No generic name can be duplicated in zoology, and one
once void for any reason cannot be revived and used in any connec-
tion. The same specific name cannot be used twice in the same
genus.

The Actual Classification of Birds has undergone radical
modification of late years, though the same machinery is employed
for its expression. This is as would be expected, seeing how pro-
foundly the theory of Evolution has affected our principles of classi-
fication, how completely the morphological has replaced other
systems, and how steadily our knowledge of the structure of birds,
and their chronological relations, has progressed. Nevertheless, the
ornithological system is still in a transition state.

With this glance at some taxonomic principles and practices, I
pass to an outline of the structure of birds, some knowledge of
which is indispensable to any appreciation of ornithological defini-
tions and descriptions. It is necessary to be brief, and I shall
confine myself mainly to the consideration of those points, and
the explanation of those technical terms, which the student needs
to understand in .order to use any systematic treatise easily and
successfully.

§ 3.— DEFINITIONS AND DESCEIPTIONS OF THE
EXTEBIOE PAETS OF BIEDS

a. Of the Feathers, or Plumage

Feathers are possessed only by birds, and all birds possess
them. Feathers are modified scales j like scales, hair, horns, plates,
sheaths, etc., they are outgrowths of the integument, or skin cover-
ing the body, and therefore belong to the class of epidermic (Gr. Itt/,
ep% upon ; derma, skin), or exoskeletal (Gr. If, ex, out ; o-kcA-

€Tov, skeleton, dried ; in the sense of “ outer skeleton ”) structures.

124

GENERAL ORNITHOLOGY

PART II

The horny coverings of the beak and feet are of the same class,
but very differently developed. Besides being the most highly
developed or complexly specialised, wonderfully beautiful and per-
fect kind of tegumentary outgrowths — besides fulfilling in a singular
manner the design of covering and protecting the body — feathers
have their particular locomotory office : that of accomplishing the act
of flying in a manner peculiar to birds. For all vertebrates, ex-
cepting birds, that progress through the air — the flying-fish, with its
enlarged pectoral fins ; the flying-reptile, with its skinny parachute ;
the flying mammal (bat) with its great webbed fingers — accomplish
aerial locomotion by means of tegumentary expansions. Birds alone
fly with tegumentary outgrowths, or appendages. All a bird’s
feathers, of whatever kind, collectively constitute its ptilosis (Gr.
TTTtAov, ptilon, a feather) or plumage (Lat. pluma, a plume).

Development of Feathers. — In a manner analogous to that of
hair, a feather grows in a little pit or pouch formed by inversion
of the dermal or true-skin layer of the integument, being formed in
a closed follicle or shut sac consisting of an inner and outer coat
separated by a layer of fine granular substance. The outer layer
or outer follicle is composed of several thin strata of nucleated
epithelial cells (cuticle cells) ; the inner is thicker, spongy, and filled
with gelatinous fluid ; a little artery and vein furnish the blood
circulation, very active during the formation of feathers. The
inner is the true matrix or mould upon which the feather is formed,
evolving from the blood-supply the gelatinous material, and resolv-
ing this into cell -nuclei; the granular layer is the formative
material which becomes the feather. The outer grows a little
beyond the cutaneous sac that holds it, and opens at the end ; from
this orifice the future feather protrudes, sprouting as a little fine-
rayed pencil point. The process is thus graphically illustrated by
Huxley : “The integument of birds is always provided with horny
appendages, which result from the conversion into horn of the cells
of the outer layer of the epidermis. But the majority of these
appendages, which are termed ‘ feathers,’ do not take the form of
mere plates developed upon the surface of the skin, but are evolved
within sacs from the surfaces of conical papillae of the dermis. The
external surface of the dermal papilla, whence a feather is to be
developed, is provided upon its dorsal [upper] surface with a median
groove, which becomes shallower towards the apex of the papilla.
From this median groove lateral furrows proceed at an open angle,
and passing round upon the under surface of the papilla, become
shallower, until, in the middle line, opposite the dorsal median
groove, they become obsolete. Minor grooves run at right angles
to the lateral furrows. Hence the surface of the papilla has the
character of a kind of mould, and if it were repeatedly dipped in

SEC.' Ill

EXTERIOR FARTS OF BIRDS

125

such a substance as a solution of gelatine, and withdrawn to cool,
until its whole surface was covered with an even coat of that sub-
stance, it is clear that the gelatinous coat would be thickest at the
basal or anterior end of the median groove, at the median ends of
the lateral furrows, and at those ends of the minor grooves which
open into them ; while it would be very thin at the apices of the
median and lateral grooves, and between the ends of the minor
grooves. If, therefore, the hollow cone of gelatine, removed from
its mould, were stretched from within ; or if its thinnest parts be-
came weak by drying ; it would tend to give way, along the inferior
median line, opposite the rod-like cast of the dorsal median groove
and between the ends of the
casts of the lateral furrows,
as well as between each of
the minor grooves, and the
hollow cone would expand
into a flat, feather-like struc-
ture with a median shaft, as
a ‘ vane ’ formed of ‘ barbs ’
and ‘barbules.’ In point of
fact, in the development of
a feather such a cast of the
dermal papilla is formed,
though not in gelatine, but
in the horny epidermic layer
developed upon the mould,
and, as this is thrust outward,
it opens out in the manner
just described. After a cer-
tain period of growth the
papilla of the feather ceases
to be grooved, and a continu-
ous horny cylinder is formed,

which C0Tiqtitlite<5 the‘nnill ’ ” Fig.19.— A partly pennaceous, partly plumulaceous

wnicn COnstlLUreS me quin. feather from Argus pheasant ; after Nitzsch. ad,

(Introd. Classif. Anim. p. 71.) calamus; a, rhachls; c, c, c, vanes,

' ^ cut a'way on left side in order not to interfere 'with &,

StPUetUPe or Feathers. the after-shaft, the -whole of the right vane of which

A perfect feather, possessing likewise cut away.

all the parts it can develop, consists of a main stem, shaft or scape (Lat.
sca][)us, a stalk ; Fig. 19, ad), and a supplementary stem or after-shaft
(hyporhachis ; Gr. vtto, hupo, under, payt?, rhachis, a spine or ridge ;
Fig. 1 9, h), each bearing two webs or vanes (Lat. vexillum, pi. vexilla,
a banner ; Fig. 19, c, c, c), one on either side. The whole scape is
divided into two parts : one, nearest the body of the bird, the tube
or barrel or “ quill ” proper (Lat. calamus, a reed), which is a hard,
horny, hollow, and semi-transparent cylinder, containing a little pith

126

GENERAL ORNITHOLOGY

PART II

Fig. 20. — Two barbs, i

in the interior ; it bears no webs. One end of this quill tapers to
be inserted into the skin j the other passes, at
a point marked by a little pit (Lat. umbilicus,
the navel) into the shaft proper or rhachis, the
second part of the stem. The rhachis is a
four-sided prism, squarish in transverse Sec-
tion, and tapers gradually to a fine point ; it is
less horny than the barrel, very elastic, opaque,
and pithy; it bears the vexilla. The after-
shaft, when well developed, is like a duplicate
in miniature of the main feather, from the
stem of which it springs, at junction of calamus
with rhachis, close by the umbilicus. It is
generally very small compared with the main
part of the feather, though quite as large in
a few kinds of birds ; it is entirely wanting
some groups of birds; it is never developed
of a vane, bearing anterior’ on the large, strong wiiig- aiid tail -feathers.

&, &, and posterior, c, barb- • , c • -in

ules; enlarged; after ihe vauc consists 01 a series ot appressed, flat,
narrowly linear or lance-linear laminae or plates,
set obliquely on the rhachis by their bases, diverging out from it at
a varying open angle, ending in a free point ; each such
narrow, acute plate is called a barb (Lat. barba, a beard ; f!

Fig. 20, a, a). Now if these laminae or barbs simply
lay alongside one another, like the leaves of a book,
the feather would have no consistency ; therefore, they
are connected together ; for, just as the rhachis bears
its vane or series of barbs, so does each barb bear its
vanes of the second order, or little vanes, called barbules
(dimin. of barba; Fig. 20, b, b, c). These are to the
barbs exactly what the barbs are to the shaft, and are
similarly given off* from both sides of the upper edges
of the barbs ; they make the vane truly a web, that is,
they so connect the barbs together that some little
force is required to pull them apart. Barbules are
variously shaped, but generally flat sideways, with
upper and lower border at base, rapidly tapering to a
slender thready end, and are long enough to reach over
several barbules of the next barb, crossing the latter
obliquely. All the foregoing structures are seen by
the naked eye or with a simple pocket lens, but the singi^ barbui^
next to be described require a microscope : they are the S^nd hoS-
barbicels (another dimin. of barba), also called cilia, or “ffgp

lashes (Fig. 21) ; and hamuli, or booklets (Lat. hamulus, Nitzsch.
a little hook; Fig. 21). These are simply a sort of fringe to the barb-

SEC. Ill

EXTERIOR PARTS OP BIRDS

127

ules, just as if the lower edge of the barbules were frayed out, and
only differ from each other in that barbicels are plain hair-like pro-
cesses, while hamuli are hooked at the end ; they are not found on
all feathers, nor on all parts of some feathers. Barbi-
cels occur on both anterior and posterior rows of
barbules, though rarely on the latter \ booklets are
confined to any anterior series of barbules, which, as
we have seen, overlie the posterior rows, forming a
diagonal mesh -work. The design of this beautiful
structure is evident; the barbules are interlocked, and
the whole made a web ; for each booklet of one barbule
catches hold of a barbule from the next barb in front,
any barbule thus holding on to as many of the barb-
ules of the next barb as it has booklets ; while, to
facilitate this interlocking, the barbules have a
thickened upper edge of the right size for the book-
lets to grasp. The arrangement is shown in Fig. 22,
where a, «, a, are four barbs in transverse section,
viewed from the cut surfaces, with their anterior,
h, h, b, b, and posterior, c, c, c, c, barbules, the former
bearing the booklets which catch over the edge of the
latter.

Types of Feathery Structure. — But all feathers
do not answer the above description. The after-shaft
may be wanting. Hooklets may not be developed,
as frequently happens. Barbicels may be few or entirely want-
ing. Barbules may be similarly deficient, or so defective as
to be only recognised by their position and relations. Even
, barbs themselves may be few or lacking on one side of the
shaft, or on both sides, as in certain bristly or hair-like styles of
feathers. Consideration of these and other modifications of feather-

Fig. 22.— Four
barbs in cross
section, a, a, a, a,
bearing anterior,
&, 6, &, b, and
posterior, c, c, c, c,
barbules, the
former bearing
hooklets which
catch over the
latter ; magni-
fied ; after
Nitzsch.

Fig. 23. — A feather from the tail of a kingbird, Tyrannus carolinensis, almost entirely
pennaceous ; no after-shaft. From nature, by Cones.

structure has led me to the recognition of three types or plans : 1.
The perfectly feathery, plumous, or pennaceous (Lat. pluma, a plume,
or penna, a feather fit for writing with ; Fig. 23), as above described.
2. The downy or plumulaceous (Lat. plumula^ a little plume, a down-
feather), when the stem is short and weak, with soft rhachis and
barbs, with long slender thready barbules, little knotty dilatations in

128

GENERAL ORNITHOLOGY

PART II

place of barbicels, and no booklets. 3. The hairy, bristly, or filoplum-
aceous (Lat. filum^ a thread), with a very long, slender stem, and rudi-
mentary or very small vanes composed of fine cylindrical barbs and
barbules, if any, and no barbicels, knots, or booklets. There is no
abrupt definition between these types of structure ; in fact, the same
feather may be constructed on more than one of these plans, as in
Fig. 19, which is partly pennaceous, partly plumulaceous. All
feathers are built upon one or another, or some combination, or
modification, of these types ; and, in all their endless diversity, may
be reduced to four or five

Different Kinds of Feathers. — 1. Contour -feathers, pennce or
plumce proper, have a perfect stem composed of calamus and rhachis,
with vanes of pennaceous structure, at least in part, usually plumu-
laceous toward the base. These form the great bulk of the surface-
plumage exposed to light; their beautiful tints give the bird’s
colours ; they are the most modified in detail of all, from the fish-like
scales of a penguin’s wings to the glittering jewels of the humming-
bird, and the endless array of the tufts, crests, ruffs, and other
ornaments of the feathered tribes ; even the imperfect bristle-like
feathers above mentioned may belong among them. Another
feature is, that they are usually individually moved by subcutaneous
muscles, of which there may be several to one feather, passing to
be attached to the sheath of the tube, inside the skin, in which the
stem is inserted. These muscles may be plainly seen under the
skin of a goose, and every one has observed their operation when a
hen shakes herself after a sand-bath, or any bird erects its top-knot.
2. Down-feathers, plumulce, are characterised by a downy structure
throughout. They more or less completely invest the body, but
are almost always hidden beneath -the contour-feathers, like pad-
ding about the bases of the latter ; occasionally they come to
light, as in the fleecy ruff about the neck of the condor, and then
usually replace contour-feathers ; they have an after-shaft, or none ;
and sometimes no rhachis at all, the barbs then being sessile in a
tuft at the end of the quill. They often stand in a regular quin-
cunx • ') between four contour-feathers. 3. Semiplumes, semi-
plumoe, may be said to unite the characters of the last two, possess-
ing the pennaceous stem of the former, and the plumulaceous vanes
of the latter; they are with or without after-shaft. They stand
among pennse, as the plumulse do, about the edges of patches of the
former, or in parcels by themselves, but are always covered by
contour-feathers. 4. Filoplumes, filoplumce, or thread-feathers, have
an extremely slender, almost invisible stem, not well distinguished
into barrel and shaft, and usually no vane, unless a terminal tuft of
barbs may be held for such. Long as they are, they are usually
hidden by the contour-feathers, close to which they stand as access-

SEC. Ill

EXTERIOR FARTS OF BIRDS

29

ories, one or more seeming to issue out of the very sacs in which
the larger feathers are implanted. These are the nearest approach
to hairs that birds have \ they are very well shown on domestic
poultry, being what a good cook finds it necessary to singe off after
plucking a fowl for the table. 5. Certain down-feathers are remark-
able for continuing to grow indefinitely, and with this unlimited
growth is associated a continual breaking down of the ends of
the barbs. Such plumulae, from being always dusted over with dry,
scurfy exfoliation, are called ;powder-clown ; they may be entitled to
rank as a fifth kind. I call them pulviplumes. They occur in the
hawk, parrot, and gallinaceous tribes, and especially in the herons
and their allies. They are always present in the latter, where they
may be readily seen as at least two large patches of greasy or dusty,
whitish feathers, matted over the hips and on the breast.

Feather Oil Gland. — Birds do not perspire, and cutaneous glands,
corresponding to the sweat-glands and sebaceous follicles so common
in Mammalia, are little known among them. But their “ oil-can ”
is a kind of sebaceous follicle, which may be noticed here in connec-
tion with other tegumentary appendages. This is a two-lobed or
rather heart-shaped gland, saddled upon the “ pope’s-nose,” at the
root of the tail, and hence sometimes called the uropygial gland
(Lat. tiropygium, rump), or rump-gland. I have named it the elceo-
dochon { Gr. lAaioSoyo?, elaiodochos, containing oil ; Fig. 24, 9). It is
composed of numerous slender tubes or follicles which secrete the
greasy fluid, the ducts of which, uniting successively in larger tubes,
finally open by one or more pores, commonly upon a little nipple-
like elevation. Birds press out a drop of oil with the beak and
dress the feathers with it, in the well-known operation called
“ preening.” The gland is large and always present in aquatic
birds, which have need of waterproof plumage ; smaller in land-
birds, as a rule, and wanting in some. The presence or absence of
this singular structure, and whether or not it is surmounted by a
particular circlet of feathers, distinguishes certain groups of birds,
and has come to be much used in classification.

Pterylogpaphy. — Feathered Tracts and Unfeathered Spaces.
— Excepting certain birds having obviously naked spaces, as about the
head or feet, all would be taken to be fully feathered. So they are
covered with feathers, but it does not follow that feathers are every-
where implanted upon the skin. On the contrary, a uniform and
continuous pterylosis is the rarest of all kinds of feathering ; though
such occurs, almost or quite perfectly, among certain birds, as the
ostrich tribe, penguins, and toucans. If we compare a bird’s skin
to a well-kept park, part woodland, part lawn ; then where feathers
grow is the woodland, where they do not grow is the lawn. The
former places are called tracts or pterylce (Gr. Trrepov, pteron, a plume
K

GENERAL ORNITHOLOGY

PART II

and v'kr]^ hyle, a wood) ; the latter, spaces or apteria (Gr. a privative,
and Ttrepov) ; they mutually distinguish certain definite areas. Not
only are the pterylce and apteria thus definite, hut their size, form,
and arrangement mark whole families and even orders of birds ; so
that pterylosis, or the formation of the feather-tracts, becomes avail-
able, and is indeed found to be important, for purposes of classifi-
cation. Pterylography, or the description of this matter, has been
made a special study by the celebrated Nitzsch, who has laid down
the general plan of pterylosis which obtains in the great majority
of birds, as follows : 1. The spinal or dorsal tract {pteryla spinalis ;
Fig. 24, 1), running along the middle of the bird above from the
nape of the neck to the tail ; subject to great variation in width, to
dilation and contraction, to forking, to sending out branches, to inter-
ruption, etc. 2. The humeral tracts {pt. humeralis ; Lat. humerus^

hand lower, surface. 1, spinal tract ; 2, humeral ; 3, femoral ; 4, capital ; 5, alar ; 6, caudal;
7, crural ; 8, ventral; 9, elgeodochon ; 10, anus.

the shoulder, or upper-arm bone ; Fig. 24, 2), always present, one
on each wing ; they are narrow bands, running from the shoulder
obliquely backward upon the upper-arm bone, parallel with the
shoulder-blade. 3. The femoral tracts {pt. femorales ; Lat. femur ^
the thigh ; Fig. 24, 3) ; a similar oblique band upon the outside of
each thigh, but subject to great variation. 4. The ventral tract
{pt. ventralis ; Lat. venter, the belly ; Fig. 24, 8), which forms most
of the plumage on the under part of a bird, commencing at or near
the throat, and continued to the vent ; like the dorsal tract, it is
very variable, is usually bifurcate, or forked into right or left
halves, with a median apterium, is broad or narrow, branched, etc.;
thus, Nitzsch enumerates seventeen distinct modifications. The
foregoing are mostly isolated tracts, that is, bands nearly surrounded
by complementary apteria ; the following are, in general, continu-
ously and uniformly feathered, and thus practically equivalent to

SEC. Ill

EXTERIOR FARTS OF BIRDS

131

the part of the body they represent : Thus, 5, the head tract {j^t.
capitalis ; Lat. ca^put, capitis^ head; Fig. 24, 4) clothes the head, and
generally runs into the beginning of both dorsal and ventral tracts.
6. The wing tract {pt. alaris ; Lat. ala, wing ; Fig. 24, 5) repre-
sents all the feathers that grow upon the wing, excepting those of
the humeral tract. 7. The tail tract {pt. caudalis ; Lat. cauda, tail ;
Fig. 24, 6) includes the tail-feathers proper and their coverts, and
those about the elseodochon, and usually receives the termination of
the dorsal, ventral, and femoral tracts. 8. The leg tract {pt.
cruralis ; Lat. crus, cruris, leg; Fig. 24, 7) clothes the legs as far as
these are feathered, which is generally to the heel, always below
the knee, and sometimes to the toes or even the claws. — I need not
enumerate the apteria, as these are merely the complements of the
pterylse. The highly important special “flight-feathers” of the
wings and “ rudder-feathers ” of the tail are to be examined beyond,
in describing those members.

Endysis and Eedysis. — Putting on and off Plumage. — Newly
hatched birds are covered for some time with a kind of down,
entirely different from such feathers as they ultimately acquire.
It is scanty, leaving much or all of the body naked, in most altricial
birds, such as are reared by the parents in the nest (Lat. altrix,
female nourisher) ; but thick and puffy in some Altrices, and in all
Prcecoces (Lat. prcecox, precocious) which run about at birth. Since
many birds which require to be reared in the nest are also hatched
clothed, or very speedily become downy, a more exact distinction
may be drawn by using the terms ptilopcedic and psilopcedic (Gr.
tttlXov, ptilon, a feather ; i/'iAo?, psilos, bare ; and irah, pais, a child)
respectively for those birds which are hatched feathered or naked
— a chicken and a canary-bird are familiar examples. It is the
rule that the higher birds are born helpless and naked, requiring to
be reared in the nest till their feathers grow ; the reverse with
lower birds, as the walking, wading, and swimming kinds. It
offers, however, many exceptions ; thus, no birds are more naked
and helpless at birth than young cormorants. Probably all prse-
cocial birds are also j^tilopsedic, and all psilopsedic birds altricial ;
but the converse is far from holding good, many Altrices, as hawks
and owls, being also ptilopsedic. In other words, psilopsedic birds
are always altricial, but ptilopsedic birds may be either altricial or
praecocial. In any case, true feathers are soon gained, in some
days or weeks — those of the wings and tail being usually the first
to sprout. The acquisition of plumage is called endysis (Gr. eVSuo-i?,
endusis, putting on). The renewal of plumage is a process familiar to
all, in its generalities, under the term “ moult,” or eedysis (Gr. eKSvens,
elcdusis, putting off). Feathers are of such rapid growth, and make
such a drain upon the vital energies, that we easily understand how

132

GENERAL ORNITHOLOGY

PART II

critical are periods of the change. The first plumage is usually
worn hut a short time ; then another more or less complete change
commonly occurs. The moult is as a rule annual ; and in many
cases more than one moult is required before the bird attains the
perfection of maturity in its feathering. It is well known how
different many birds are the first year in their coloration from
that afterward acquired. Sometimes changes progress for several
years; and some birds appear to have a period of senile decline.
All such changes are necessarily connected, if not with actual
moult, as is the rule, then at any rate with wear and tear and
repair of the plumage. The first plumage being gained, under
whatever conditions peculiar to the species, it is the general rule
that birds are subject to single or annual moult. This commonly
occurs in the fall, when the duties of incubation are concluded, and
the well-worn plumage most needs renewal. Many, however, moult
twice a year, the additional moult usually occurring in the spring-
time, when a fresh nuptial suit is acquired ; in such cases the moult
is said to be double or semi-annual. Such additional moult is
generally incomplete ; that is, all the feathers are not shed and re-
newed, but more or fewer new ones are gained, with more or less
loss of the old ones, if any. The most striking ornaments donned
for the breeding season, as the elegant plumes of many herons, are
usually worn but a brief time, being doffed in advance of the
general fall moult. A few birds, as the ptarmigan (Lagopus),
regularly have even a third or triple moult, shedding many of their
feathers as usual in the early autumn, then changing entirely to
pure white for the winter, then in spring moulting completely to
assume their wedding-dress. As a rule, feathers are moulted so
gradually, particularly those of the wings and tail, and so simultane-
ously upon right and left sides of the body, that birds are at no
time deprived of the power of flight. The first flight -feathers
acquired by young birds are usually kept till the next season ; but
in those that fly very early, before they are half-grown, as so many
gallinaceous birds do, their first weak wing-feathers are included in
the general moult which occurs to young and old in the fall. The
duck tribe offer the remarkable case that they drop their wing-
quills so nearly all at once as to be for some time deprived of the
power of flight. It is quite certain that many birds change the
colours of their plumage remarkably, without losing or gaining any
feathers, by some process which affects the texture of the feathers,
such as the shedding of the barbicels and booklets, or its pigmenta-
tion, or by such processes combined. The male bobolink {Dolichonyx
oryzivorus) changes from the buff dress of the female to his rich black
suit without losing or gaining any feathers, a process which is called
aptosochromatism. It is difiicult to lay down any rules of moulting

SEC. Ill

EXTERIOR FARTS OF BIRDS

133

for particular groups of birds, since birds very closely related differ
greatly in respect to their changes of plumage \ and the subject
has not yet received the attention its interest and importance
should claim for it. The physiological processes involved are
analogous to those concerned in the shedding of the hair of
mammals and the casting of the cuticle of reptiles.

Plumage-changes with Sex, Age, and Season. — Aside from
any consideration of the way in which plumage changes, whether
by moult or otherwise, the fact remains that most birds of the same
species differ more or less from one another according to certain
circumstances. The dissimilarity is not only in coloration, though
this is the usual and most pronounced difference, but also in the
degree of the development of plumes — their size, form, and texture.
Since young birds are those which have not come to sexual vigour ;
since breeding recurs at regular periods of the year ; and since
males and females usually differ in plumage, nearly all the various
dresses worn by different individuals of the same species are cor-
related with the conditions of the reproductive system. As the
internal generative organs represent of course the essential or
'primary sexual characters,' all those features of the plumage just
indicated may be properly classed as secondary sexual characters.
These are of great importance, not only in practical ornithology,
but as the basis of some of the soundest views that have been
advanced respecting the evolution of specific characters in this class
of animals. The generalisations may be made : that when the
sexes are strikingly different in plumage, the young at first resemble
the female ; when the adults are alike, the young are different from
either ; when seasonal changes are great, the young resemble the
fall plumage of the parents ; and further, that when the adults of
two related species of the same genus are nearly alike, the young
are usually intermediate, their specific characters not being fully
developed. Specific characters are often to be found only in the
male, the females of two related species being scarcely distinguish-
able, though the males may be told apart at a glance. Extraordinary
developments of feathers, as to size, shape, and colour, are often
confined to one sex, usually the male. The more richly, exten-
sively, or peculiarly the male is adorned, the simpler the female in
comparison, as the peacock and peahen. Darwin has formulated
the several categories of secondary sexual characters, giving the
following rules or classes of cases : “1. When the adult male is
more beautiful or conspicuous than the adult female, the young of
both sexes in their first plumage closely resemble the adult female,
as with the common fowl and peacock, or, as occasionally occurs,
they resemble her much more closely than they do the adult male.
2. When the adult female is more conspicuous than the adult male.

134

GENERAL ORNITHOLOGY

PART II

as sometimes, though rarely, occurs [chiefly with certain birds of
prey and snipe-like birds], the young of both sexes in their first
plumage resemble the adult male. 3. When the adult male
resembles the adult female, the young of both sexes have a peculiar
first plumage of their own, as with the robin [usual]. 4. When the
adult male resembles the adult female, the young of both sexes in
their first plumage resemble the adults [unusual]. 5. When the
adults of both sexes have a distinct winter and summer plumage,
whether or not the male differs from the female, the young resemble
the adults of both sexes in their winter dress, or much more rarely
in their summer dress, or they resemble the females alone. Or the
young may have an intermediate character; or again, they may
differ greatly from the adults in both their seasonal plumages. 6.
In some few cases the young in their first plumage differ from each
other according to sex ; the young males resembling more or less
closely the adult males, and the young females more or less closely
the adult females.” — {Desc. of Man, ed. 1881, p. 466.)

h. The Topography of Birds.

The Contour of a Bird with the feathers on is spindle-shaped
or fusiform (Lat. fusus, a spindle), tapering at both ends ; it
represents two cones joined base to base at the middle or greatest
girth of the body, tapering in front to the tip of the bill, behind to
the end of the tail. The obvious design is easiest cleavage of air
in front, and least drag or wash behind, in the act of flying. This
shape is largely produced by the lay of the plumage ; a naked bird
presents several prominences and depressions, this irregular contour
being reducible, in general terms, to two spindles or double cones.
The head tapers to a point in front, at the tip of the bill, and
contracts behind, toward the middle of the neck, in consequence of
diminution in bulk of the muscles by which it is slung on the neck ;
which last is somewhat contracted or hour-glass-shaped near the
middle, swelling where it is slung to the body. The body is largest
in front and tapers to the tail. The

Centre of Gravity is admirably preserved beneath the centre
of the body, and opposite the points where it is supported by the
wings. The enormous breast - muscles of a bird are among its
heaviest parts, sometimes weighing, to speak roundly, as much as
one-sixth of the whole bird. 'Now these are they that effect all the
movements of the wings at the shoulder-joints, lifting as well as
lowering the wings. Did these pectoral muscles pull straight, the
lifters of the wing would have to be above the shoulder- joint ; but
they all lie below it, and the lifters accomplish their ofiice by
running through pulleys to change the line of their traction. They

SEC. Ill

EXTERIOR FARTS OE BIRDS

135

work like men hoisting sails from the deck of a vessel ; and thus,
like a ship’s cargo, a bird’s chief weight is kept below the centre of
motion. Top-heaviness is further obviated by the way in which
birds with a long heavy neck aud head draw these parts in upon
the breast, and extend the legs behind, as is well shown by the
attitude of a heron flying. The nice adjustment of balance by the
variable extension of the head and feet is exactly like that produced
in weighing by shifting a weight along the arm of a steelyard ;
and together with the slinging of the chief weight under the wings
instead of over or even between them, enables a bird to easily keep
right side up in flight. The

Exterior of a Bird is divided for purposes of description into
seven — 1. The head (Lat. caput)-, 2. The neck (Lat. collum) ;
3. The body proper, or trunk (Lat. truncus) ; 4. The bill or beak
(Lat. rostrum) ; 5. The wings (Lat. pi. alee) ; 6. The tail (Lat. cauda)-,
7. The feet (Lat. pi. pedes). Of these, 1, 2, 3, the head, neck, and
trunk, are collectively termed the body (Lat. corpus), in distinction
from 4, 5, 6, 7, which are the members (Lat. pi. membra). The
wings and feet are of course double or paired parts. The bill is
strictly but a part of the head ; but its manifold uses as an organ
of prehension make it functionally a hand, and therefore one of the
“ members.” The

Head has the general shape of a four-sided pyramid ; of which
the base is applied to the end of the neck, therefore not appearing
from the exterior, and the apex of which is frustrated at the base
of the bill. The uppermost side is more or less convex or vaulted,
sloping in every direction ; the under side is flattish and horizontal ;
the lateral surfaces are flattish and vertical ; all similarly taper
forward. The departures from any such typical shape are endless
in degree and variable in kind, giving rise to numerous general
descriptive terms, such as “head flattened,” “head globular,” but
not susceptible of exact deflnitiou. The head is moulded, of course,
upon the skull, corresponding in a general way to the brain-cavity
of the cranium proper, both in size and shape but it differs in
several particulars. In the first j)lace, there is the scaffolding of
the jaws ; secondly, large excavations to receive the eyeballs, and
smaller ones for the ear-parts ; thirdly, muscular masses overlying
the bone ; and lastly, in some birds, large hollow spaces in the bone
between the inner and outer tables or plates of the cranial walls.
Each side of the head presents two openings for the eye (Lat. oculus)
and ear (Lat. auris), the position of which is variable, both absolutely
and in relation to each other. But in the vast majority of birds,
the eye is strictly lateral in situation, and near the middle of the
side of the head ; while the ear is behind and a little below the
eye, near the articulation of the lower jaw. But the shape of the

136

GENERAL ORNITHOLOGY

PART II

skull of owls is such, that the eyes are directed forward, and such
birds are said to have “eyes anterior.” Owls also have enormous
outer ears, in some cases provided with a movable flap or conch,
closing upon the opening like the lid of a box ; in many cases the
ear-parts, and some of the cranium itself, being unsymmetrical. In
most birds the ear-opening is quite small, and only covered by
modified feathers, the ear-coverts or auriculars. In the woodcock
and snipe, owing to the way the brain-box is tilted up, the ears are
below and not behind the eyes. The mouth (Lat. os, gen. oris) is
always a fissure across the front of the head. The cleavage varies,
both in extent and direction ; the latter is usually horizontal, or
nearly so, but may trend much downward ; the former varies from
a minimum, in which the cleft does not reach back of the horny
part of the bill, as in a snipe, to the maximum seen in fissure-billed
birds like the swifts and goatsuckers, which gape almost from ear
to ear. There are no other openings in the head proper, for the
nostrils are always in the bill. The

Neek, in effect, is a simple cylinder, rendered somewhat hour-
glass-shaped, as above said. It consists of a movable chain of
bones, the cervical vertebrce (Lat. cei^vix, the neck ; verto, I turn)
enveloped in muscle, along which in front lie the gullet (Lat.
oesophagus) and windpipe (Lat. trachea), with associate blood-vessels,
nerves, etc. Its length is very variable, as is the number of its
bones, the latter ranging from 8 to about 26. Bearing as it does
the head, with the hill, which is the true hand of a bird, the neck
is extremely flexible, to permit the necessarily varied movements
of this handy member. Its least length may be said to be that
which allows the point of a bird’s beak to reach the oil-gland on
the rump ; its greatest length sometimes exceeds that of the body
and tail together, as in the case of a swan, crane, or heron. The
length is usually in direct proportion to that of the legs, in obvious
design of allowing the beak to touch the ground easily to pick up
food. The neck is habitually carried in a double curve, like an
open S or italic/, the lower belly of the curve, convex forward,
fitting in between the forks of the merrythought (Lat. furculum),
the upper curve holding the head horizontal at the same time.
This “ sigmoid flexure ” {sigma, Greek S), highly characteristic of
the bird’s neck, is produced hy the saddle-shaping of the articular
surfaces of the several bones. The mechanical arrangement is such,
that the sigma may be easily bent till the upper end (head) rests on
the lower convexity, or as easily straightened to a right line ; but
little if any further deviation in opposite curvature is permitted.
As a generalisation, the neck may be called relatively longest in
wading birds, as herons, cranes, ibises, etc. ; shortest in perching
birds, as the great majority of small Fasseres ; intermediate in

SEC. Ill

EXTERIOR TARTS OF BIRDS

137

swimming birds. But many swimmers, as swans and cormorants,
have extremely long necks ; and some waders, as plovers, have
very short ones. A long neck is a rarity among the higher birds
(above the Gallince), in most of which the head seems to nestle upon
the shoulders. The longer the neck, the more sinuous and flexible
is it likely to be. Anatomically, the neck ends in front at the
articulation of the atlas (first cervical vertebra) with the skull, and
behind at the first vertebra which bears free jointed ribs reaching
the sternum. (See also § 4, Anatomy.) The shape of the

Body proper, or Trunk, is obviously referable to that of the
egg; it is ovate (Lat. ovum, an egg; whence oval, the plane figure
represented by the middle lengthwise section of an egg ; ovate or
ovoid, the solid figure). The swelling of the breast represents the
greatest diameter of the egg, usually near the larger end. But the
ovoid is never perfectly expressed, and departures from the figure
are numberless. In general, the higher perching birds have the
body nearly of the ovate shape ; among waders, the figure is usually
compressed, or flattened vertically, as is well seen in the herons, and
still better in the rails, where the lateral narrowing is at an extreme ;
among swimmers, the body is always more or less depressed, or
flattened horizontally, and especially underneath, that the birds
may rest on the water with more stability, as well shown by a duck
or diver. Anatomically the body begins with the foremost dorsal
or thoracic vertelrce, or those that bear true ribs ; laterally, it ceases
quite definitely at the shoulder-joints, the whole of the fore limb
being outside the general content of the trunk; behind, in the
middle line, it includes everything, only the t^dl-feathers themselves
being beyond it ; behind and laterally, it includes more or less of
the legs, for these are generally buried in the common integument
of the body to the knee-joint, nearly or quite so, and sometimes to
the heel-joint ; though in anatomical strictness the trunk should be
limited by the hip-joint. The rib-bearing part of the back-bone,
the ribs themselves, and the greatly enlarged breast-bone (Lat.
sternum) compose the walls of the chest (Lat. thorax). Upon this
bony box, which contains the heart and lungs and some other
viscera, are saddled on each side the bones of the shoulder-girdle or
scapular arch, namely, the shoulder-blades (Lat. scapulce), the coracoids,
and the collar-bones (Lat. claviculce), all three of which, on each side,
come together at the shoulder-joint (Figs. 1, 2). The thoracic cavity is
not separated by any partition or diaphragm from that of the belly
(Lat. abdomen), which with the pelvis, or basin, contains the digestive,
urinary, and genital organs. The pelvis is composed, in dorsal
mid-line, of so many of the vertebrae {dorso-lumbar, sacral proper, and
urosacral), as become immovably joined to one another, and later-
ally of the confluent haunch - bones. The numerous anchylosed

GENERAL ORNITHOLOGY

PART II

138

(or confluent) vertebrae compose the sacrum. The haunch-bones or ossa
innominata consist on each side of three bones, ilium, ischium, and
pubis, in adult life more or less perfectly anchylosed. Where they all
three come together is the hip-joint. The remaining bones, usually
included among those of the body proper, are the coccygeal or caudal
vertebrae. (For anatomical details see beyond, under Osteology, etc.)

Topography of the Body. — Besides being thus' divided into
head, neck, trunk, and members, the exterior of the body is further
subdivided or mapped out into regions for the purposes of description.
It is necessary for the student to become familiar with the “ topo-
graphy ” of a bird, as this kind of mapping out may be called, for
the names of the regions or outer areas are incessantly used in
ordinary descriptive ornithology. Many more names have been
applied than are in common use ; I shall try to define and explain
all those which are usually employed, beginning with the parts of
the body, and ending with those of the members.

1. Regions of the Body.

Upper and Under Parts. — Draw a line from the corner of the
mouth along the side of the head and neck to and through the
shoulder- joint and thence along the side of the body to the root
of the tail ; all above this line, including the upper surfaces of the
wings and tail, are upper parts; all below it, including under
surfaces of wings and tail, are under parts; for which the short
words “above” and “below” often stand. The distinction is
purely arbitrary, but so convenient as to be practically indispens-
able. It will be seen how an otherwise lengthy description,
enumerating parts that lie over or under the “ lateral line,” can be
put in so few words as, for example, “above, green ; below, yellow.”
Many birds’ colours have some such simple general distribution.
These parts are also the dorsal (Lat. dorsum, back) and ventral (Lat.
venter, belly) surfaces or aspects. The upper parts of the body
proper, or trunk, have also received the general name of notceum
(Dr. vwTos, notos, back) ; the under parts, similarly restricted, that
of gastrceum (Gr. yacrrrip, gaster, belly) : but these terms are not
much used. These two are never naked, while both head and neck
may be variously bare of feathers. The only exception is the
transient condition of certain birds during incubation, when, like
the eider-duck, they pull off feathers to furnish the nest, or when
the plumage, as usually happens, wears off. The gastrseum is
rarely ornamented with feathers different in texture or structure
from those of the plumage at large ; but such a case is furnished
by Lewis’s woodpecker (Asyndesmus torquatus). The notseum, on
the contrary, is often the seat of extraordinary development of

SEC. Ill

EXTERIOR TARTS OF BIRDS

139

feathers, either in size, shape, or texture, or all three of these
qualities ; as the singularly elegant dorsal plumes of many herons.
Individual feathers of the notseum are generally pennaceous, and
for the most part straight and lanceolate ; and as a whole lie
smoothly shingled or imbricated. The ventral feathers are usually
more largely plumulaceous, and less flat and imbricated, but even

coverts. 15, tail. 16, under tail-coverts (crissum). 17, tarsus. 18, abdomen. 19, hind toe
(hallux). 20, gastrceum, including 18 and 24. 21, outer or fourth toe. 22, middle or third toe.
23, side of the body. 24, breast (peciifs). 25, primaries. 26, secondaries. 27, tertiaries ; Nos.
25, 26, 27 are all remiges. 28, primary coverts. 29, alula, or bastard wing. 30, greater coverts.
31, median coverts. 32, lesser coverts. 33, the “ throat,” including 34, 37, 38. 34, jugulum or

lower throat. 35, auriculars. 36, malar region. 37, gula, or middle throat. 38, mentum, or
chin. 39, angle of commissure, or corner of mouth. 40, ramus of under mandible. 41, side
of under mandible. 42, gonys. 43, apex, or tip of bill. 44, tomia, or cutting edges of the bill.
45, culmen, or ridge of upper mandible, corresponding to gonys. 46, side of upper mandible.
47, nostril. 48 passes across the bill a little in front of its base.

more compact, that is, thicker, than those of the upper parts ;
especially among water birds, where they are more or less curly,
and very thick-set. There are subdivisions of the

Notseum. — Beginning where the neck ends, and ending where
the tail-coverts begin (see Fig. 25, 12), this part of a bird is sub-
divided into back (Lat. dorsum ; Fig. 25, 11) and rump (Lat. uropy-
gium ; Fig. 25, 13). These are in direct continuation of each other.

140

GENERAL ORNITHOLOGY

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and their limits are not precisely defined ; the feathers of both are
of the ^teryla dorsalis. In general, we should call the anterior two-
thirds or three-fourths of notseum “hack,” and the rest “rump.”
With the former are generally included the scapular or shoulder-
feathers, scapulars or scapularies ; these are they that grow on the
pterylce liumerales. The region of notseum they represent is called
the scapulare, and that part of notaeum strictly between them is
called the inUr scapular c (Fig. 25, 10); it is often marked, as in the
house sparrow, with streaks or some other distinctive coloration.
A part of dorsum, lying between interscapulare and uropygium, is
sometimes recognised as the “ lower back ” (Lat. tergum) ; but this
distinction is not practically useful. To uropygium probably also
belong the feathers of the pterylce femorales, or at any rate these are
commonly included with the rump in descriptions ; but they more
properly represent the flanks (Lat. ilia, or hypochondria) ; that is,
sides of the rump. They are sometimes the seat of largely
developed or otherwise peculiarly modified feathers, as the snowy
flank-plumes of the white-bellied swift {Panyptila saxatilis) or violet-
green swallow {Tachycineta thalassina), which meet over the rump.
The whole of notseum, taken together with the upper surfaces of
the wings, is called the mantle (Lat. stragulum, a cloak) ; often a
convenient term, as, for example, in describing gulls and terns. In
like manner, the

Gastpseum is subdivided into regions, called, in general terms,
hreast (Lsit. pectus ; Fig. 25, 24), lelly (Lat. abdomen; Fig. 25, 18),
and sides of the body (Lat. pleurce ; Fig. 25, 23). The “sides” or
pleurse belong really as much to the dorsal as to the ventral aspects
of a bird’s body ; but in consequence of the underneath-freighted
shape, the line we drew passes so high up along them, that they
are almost entirely given over to gastrseum. The breast begins over
the merrythought where jugulum (see beyond) ends ; on either
hand, it slopes up to “ sides ” ; behind, its extension is indefinite.
It should properly reach as far as the breast-bone does, to the limit
of the thorax ; but in many birds this would leave almost nothing
for abdomen, and the limit would moreover fluctuate with almost
every family of birds, the sternum being so variable in length.
Practically, therefore, without reference to the breast-bone, “ breast ”
or pectus is restricted to the swelling anterior part of gastrseum,
which we call belly or abdomen as soon as it begins to straighten
out and flatten. Abdomen, like pectus, rounds up on either hand
into sides ; behind, it ends definitely in a transverse line passing
across the anus. It has been unnecessarily divided into epigastrium
or “pit of the stomach,” and venter or lower belly; but these terms
are rarely used. (Crissum is a word constantly used for some in-
definite region immediately about the vent ; sometimes meaning

SEC. Ill

EXTERIOR TARTS OE BIRDS

141

the flank, sometimes the vent-feathers or under tail-coverts proper \
I refer to it again in connection with these last.) Though these
boundaries seem fluctuating and not perfectly satisfactory, a little
practice will enable the student to appreciate their proper use in
descriptions, and to employ them himself with sufiicient accuracy.
The adjectival terms are respectively pectoral, abdominal, and lateral.
The anterior continuation of the trunk, or the

Neck (Lat. collum) is likewise subdivided into regions. Its
lateral aspects, except in those birds that have lateral neck-tracts
of feathers, are formed by the meeting over its sides of the feathers
that grow on the dorsal and ventral pterylse, the skin being usually
not planted with feathers. Partly on this account, perhaps, a dis-
tinctively named region is not often expressed; we say simply
“sides of the neck,” or “neck laterally” [parauchenia ; Fig. 25, 9).
The neck behind, or the dorsal (upper) aspect, is divided into two
portions : a lower, the “ hind neck ” proper, or “ scruff of the neck ”
(Lat. cervix ; Fig. 25, 8), next to the back; and an upper, or “nape
of the neck” (Lat. nucha; Fig. 25, 7), adjoining the hind head.
These are otherwise respectively known as the cervical and nuchal
region; and, in speaking of both together, we usually say “the
neck behind.” The front of the neck has been needlessly sub-
divided, and these subregions vary with almost every writer. It
suffices to call it throat (Lat. gula. Fig. 25, 37, or jiigulum, 34);
remembering that the jugular portion is lowermost, vanishing in
breast, and the gular uppermost, running into chin along the under
surface of the head. Guttur is a term sometimes used to include
gula and jugulum together : it is simply equivalent to “ throat,” as
just defined ; the adjective is guttural. Though generally covered
with feathers, the neck, unlike the trunk, is frequently partly naked.
When naked behind, it is usually cervix that is bare, as so charac-
teristically occurs in herons, from interruption of the forward
extension of the pteryla spinalis. Nucha is seldom if ever naked,
except as an extension of general bald-headedness. Gula is
similarly naked from above downwards, as conspicuously illustrated
in the order Steganopodes, comprising the pelicans, cormorants, etc.,
which have a bare gular pouch ; and as seen in many vultures,
whose baldness extends over nucha and gula, and even all around
the neck, as in the condor, whose nakedness ends with so singular
a collar of close-set, downy feathers. The lower throat or jugulum
becomes naked in a few birds, in which a distended crop or craw
protrudes, pushing apart feathers of two branches of the ptenjla
ventralis as these ascend the neck. The rule is, that the neck is
not the seat of enlarged or otherwise highly developed feathers,
which might restrict the requisite freedom of its motion ; but there
are some signal exceptions, among which may be instanced the

142

GENERAL ORNITHOLOGY

PART II

grouse family. The ruffed grouse^ has a singular umhrella-like
tuft on each side of the neck : the pinnated grouse ^ has still more
curious winglets in the same situation, covering bare distensible
skin: the sharp -tailed grouse^ is in somewhat similar but less
pronounced case ; while the cock-of-the-plains ^ has some extra-
ordinary jugular developments of feathers in connection with his
subcutaneous tympanum. Cervix proper almost never has modified
feathers, but often a transverse coloration different from that of the
rest of the upper parts ; when conspicuous, this is called “ cervical
collar,” to distinguish it from the guttural or jugular “ collars ” or
rings of colour. Nucha is frequently similarly marked with a
“ nuchal band ” ; often special developments there take the form of
lengthening of the feathers, and we have a “nuchal crest.” More
particularly in birds of largely variegated colours, guttur and
jugulum are marked lengthwise with stripes and streaks, of which
those on the sides are apt to be different from those along the
middle line in front. Jugulum occasionally has lengthened feathers,
as in many herons. Higher up, the neck in front may have
variously lengthened or otherwise modified feathers. Conspicuous
among these are the ruffs, or tippets, of some birds, especially of the
grebe family {Podicipedidce), and, above all other English birds, of
the male ruff {Machetes pugnax). But these, and a few other modi-
fications of the feathers of the upper neck, are more conveniently
considered with those of the

Head. — Though smaller than any of the areas already con-
sidered, the head has been more minutely mapped out, and much
detail is required by the number and importance of its recognisable
parts or regions. Without intending to mention all that have
been named, I describe all needed to be known for any practical
purposes.

“ Top of the head ” is a collective term for all the upper surface,
from base of bill to nape, and laterally to about the level of the
upper border of the eyes ; this is the ffleum or “ cap ” (Fig. 25, 1, 4,
6) ; it is divided into three portions. The forehead, or frontal
region, or simply “the front” (Lat. frons ; Fig. 25, 1), includes all
that slopes upward from the bill, — generally to about opposite the
anterior border of the eyes. Middle head or crown (Lat. corona) or
vertex (Lat. Fig. 25, 1) includes the top of the head proper, or
highest part, from the rise of the forehead to the fall of the hind-
head towards nucha. This slope is the hind-head, or occiput (Lat.
Fig. 25, 6). The lateral border of all three constitutes the super-
ciliary line, that is, the line over the eye (Lat. super, over ; cilia,

^ Bonasa umhellus, wliich closely resembles the European hazel - grouse, B.
hetuUna. ^ Gupidonia cupido.

^ Pedioecetes phasiandlus. ^ Centrocercus urophasianus.

SEC. Ill

EXTERIOR FARTS OF BIRDS

143

little hairs, especially of the brows). ‘‘Crown” is often used as
the same thing as pileum. The adjectives of the several words are
frontal, coronal or vertical, and occipital : pileum has none in use,
coronal being said instead.

“ Side of the head ” is a general term defining itself ; it presents
for consideration several regions. The orbital or circum-orbital region,
or simply the orbit (Lat. orbis, an orb, here the socket of the eyeball ;
Fig. 25, 3), is a small space forming a ring around the eye. It
includes the eye, and especially the eyelids (Lat. palpebrce). The
points where these meet, in front and behind, respectively, are the
anterior canthus and posterior canthus (Gr. KavOos, Icanthos, Lat. canthus,
a tire). The orbital region is subdivided into supra-orbital, infra-
orbital, ante-orbital, and post-orbital, according as its upper, under,
front, or back portion is desired to be specially designated. The
situation of the orbit varies much in different groups of birds ; it is
generally midway, as said above, but may be higher or lower,
pressed on toward the bill, or pushed far up and back, as strikingly
shown in the woodcock {Scolopax rusticula). In owls, the orbital
region is exaggerated into a great disk of radiating feathers, con-
ferring a peculiar physiognomy. The aural or auricular (Lat. amis,
or auriculum, ear; Fig. 25, 35) region lies about the external
opening of the ear, or meatus auditorius ; its position varies in heads
of different shapes, but it nearly always lies behind and a little
below the eye. Wherever located, it may be recognised at a glance,
by the peculiar texture of the feathers (the auriculars) which over-
lie the meatus. Doubtless to offer least obstacle to sound, these
are a parcel of loose-webbed little plumes, which may be collectively
raised and turned forward, exposing the orifice of the ear ; they
are extremely large and notable in those owls which have com-
plicated external ear-parts, and in such they form part of the great
facial disc. The term “ temporal region ” or “ temple ” is not often
used in ornithology, not being well distinguished from the post-
orbital space between eye and ear, and having nothing special about
it. At the lowermost back corner of the side of the head, generally
just behind and below the ear, may be seen or felt a hard protuber-
ance ; it is the sharpest corner-stone of the head, being the place
where the lower jaw hinges upon the skull. This is called the
“ angle of the jaw ; ” it is a good landmark, which must by no
means be confused with the “angle of the mouth,” where the horny
parts of the beak come together. The lore (Lat. lorum, a strap, or
bridle; hence, place where the cheek-strap passes; Fig. 25, 2)
includes pretty much all the space between the eye and the side of
the base of the upper mandible ; a considerable part of it is simply
ante-orbital. Thus we say of a hawk, “lores bristly;” and ex-
amination of a bird of that kind will show how large a space is

144

GENERAL ORNITHOLOGY

PART II

covered by the term. Lore, however, should properly be restricted
to a narrow line between the eye and bill in the direction of the
nostrils. It is excellently shown in the heron and grebe families,
where “naked lores” is a distinctive character. The lore is an
important place, not only from being thus marked in many birds,
but from being frequently the seat of specially modified or specially
coloured feathers. The rest of the side of the head, including the
space between angle of jaw and bill, has the name of cheek (Lat.
gena, first eyelid, then, and generally, the prominence under the
eye formed by the cheek-bones; Fig. 25, 36). It is bounded above
by loral, infra-orbital, and auricular regions ; below, by a more or
less straight line, representing the lower edge of the bony prong of
the under mandible. It is cleft in front for a varying distance by
the backward extension of the gape of the mouth ; above this gape
is more properly gena, or malar region (Lat. mala, upper jaw) in
strictness ; below it is jaw {maxilla), or rather “ side of the jaw.”
The low^er edge of the jaw definitely separates the side of the head
from the “ under surface ” of the head ; properly bounded behind
by an imaginary line drawn straight across from one angle of the
jaw to the other, and running forward to a point between the forks
of the under mandible. As already hinted, “throat” (gula ; Fig.
25, 37) extends upward and forward into this space without
obvious dividing line ; it runs into chin (Lat. mentum ; Fig. 25, 38),
of which it is only to be said that it is the (varying in extent)
anterior part of the under surface of the head. Anteriorly, it may
be conveniently marked off, opposite the point where the feathers
end on the side of the lower jaw, from the feathery space (when
any) hehveen the branches of the upper mandible itself ; this latter
is called the interramal space (Lat. inter, between, ramus, fork).

The head is so often marked lengthwise with different colours,
apt to take such definite position, that these lines have received
special names. Median vertical line is one along the middle of pileum,
from base of bill to nucha : lateral vertical lines bound it on either
side. Superciliary line has already been noticed ; below it runs the
lateral stripe ; that part of it before the eye is loral or ante-orbital ;
behind the eye, post-orbital ; when these are continuous through the
eye, they form a transocular (Lat. trans, across ; oculus, eye) line ;
below this is malar line, or cheek-stripe (Lat. frenum, a bridle) ;
below this on the under jaw, maxillary or sulmaxillary line; in the
middle below, mental (L. mentum, chin) or gular lines.

No other part of the body has so variable a ptilosis as the head.
In the great majority of birds it is wholly and densely feathered ;
it ranges from this to wholly naked ; but nakedness, it should be
observed, means only absence of perfect feathers, for most birds
with unfeathered heads have a hair-like growth of filoplumes on the

SEC. Ill

EXTERIOR PARTS OF BIRDS

145

skin. Examples of naked - headed birds are the turkey, the
vultures, the cranes, and some of the heron tribe, as ibises.
Associated with more or less complete baldness, is the frequent
presence of various fleshy outgrowths, as com5s, wattles, caruncles,
lohes, and flaps of all sorts, even to enumerate which would exceed
our limits. The parts of the barn-yard cock exemplify the whole.
Sometimes horny plates take the place of feathers on part of the head ;
as the frontal shields of the coots and gallinules. A very common
form of head-nakedness marks one whole order of birds, the Stegano-
podes, which have mentum and more or less of gula naked and trans-
formed into a sort of pouch, extremely developed in the pelicans,
and well seen in the cormorants. The next commonest is definite
bareness of the lores, as in all herons and grebes ; in the former
including the whole circum-orbital region. A little orbital space is
bare in many birds, as the vulturine hawks, and some pigeons ;
species of grouse have a bare warty supra-orbital space. Among
water birds particularly, more or less of the interramal space is
almost always unfeathered ; the nakedness always proceeds from
before backwards.

The opposite condition, that of redundant feathering, gives rise
to all the various crests (Lat. pi. cristce) that form such striking
ornaments of many birds. Crests proper belong to the top of the
head, but may be also held to include those growths on its side ;
these together being called crests in distinction to the ruffs, ruffles,
beard, etc., of gulla or mentum. Crests may be divided into two
kinds : 1, where the feathers are simply lengthened or otherwise
enlarged; and 2, where the texture, and sometimes even the
structure, is altered. Nearly all birds possess the power of moving
and elevating the feathers on the head, simulating a slight crest in
moments of excitement. The general form of a crest is a full, soft
elongation of the coronal feathers collectively ; when perfect, such a
crest is glolular, as in the Neotropical genus Pyrocephalus ; generally,
however, the feathers lengthen on the occiput more than on the
vertex or front, and this gives us the simplest and commonest form.
Such crests, when more particularly occipital, are usually connected
with lengthening of nuchal feathers, and are likely to be of a thin,
pointed shape, as is well shown in the lapwing {Vanellus cristatus).
Coronal or vertical crests proper are apt to be rather different in
coloration than in specially marked elongation of the feathers ; they
are perfectly illustrated in the goldcrest, and other species of the
genus Regulus. Frontal crests are the most elegant of all ; they
generally rise as a pyramid from the forehead, as excellently
shown in the crested titmouse {Parus cristatus) and others. All the
foregoing crests are generally single, but sometimes double ; as shown
in the two lateral occipital tufts of the “ horned ” lark (Premophila
L

146

GENERAL ORNITHOLOGY

PART II

alpestris)^ in all the “ horned ” owls, and in a few cormorants. Lateral
crests are always double, one on each side of the head ; they
are of various shapes, but need not be particularised here, especially
since they mostly belong to the second class of crests, — those con-
sisting of texturally modified feathers. It is a general, though not
exclusive, character of these last that they are tem^pmury ; while the
other kind is only changed with the general moult, these are assumed
for a short period only, the breeding season ; and, furthermore, they
are often distinctive of sex. Occurring on the top of the head, they
furnish the most remarkable ornaments of birds. I need only
instance the elegant helmet-like plumes of the partridges of the
genus Lopliortyx ; the graceful flowing train of Oreortyx ; the some-
what similar plumes of the night-herons. The majority of the
cormorants, and many of the auks, possess lateral plumes of similar
description ; these, and those of the herons, are probably — in most
cases certainly — deciduous; while those of the partridges above
mentioned last as long as the general plumage. These lateral
plumes, in many birds, especially among grebes, are associated with,
and, in fact, coalesce with, the ruffs, which are singular lengthening
and modifying in different ways of feathers of auriculars, gense, and
gula ; and are almost always temporary. Beards, or special lengthen-
ing of the mental feathers alone, are comparatively rare ; a European
vulture, Gypa'etus barhatus, is a good example. The feathers some-
times become scaly {squamous), forming, for instance, the exquisite
gorgelets or frontlets of humming-birds. They are often bristly
{setaceous), as about the lores of nearly all hawks. A particular set
of bristles, which grow in single series along the gape of many birds,
are called rictal bristles or vibrissce. These occur in greater or less
development in most small insectivorous birds ; they are large and
stiff and highly characteristic of the family Muscicapidce or flycatchers;
while in some of the goatsuckers {Caprimulgidoe) they are pro-
digiously long, and in one species of that family {Antrostomus
carolinensis) they have lateral filaments. While usually all the un-
lengthened head-feathers point backward, they are sometimes erect
forming a velvety pile, or they may radiate in a circle from a given
point, as from the eye in most owls, where they form a facial dish

Of the Members : their Parts and Organs

I. THE BILL

The Bill (Lat. rostrum) is hand and mouth in one : the instru-
ment of prehension. As hand, it takes, holds, and carries food or
other substances, and in many instances feels ; as mouth, it tears,
cuts, or crushes, according to the nature of the substances taken ;

SEC. Ill

EXTERIOR TARTS OF BIRDS

147

assuming the functions of both lips and teeth, neither of which do
any recent birds possess. An organ thus essential to the prime
functions of birds, one directly related to their various modes of life,
is of much consequence in a taxonomic point of view ; yet its
structural modifications are so various and so variously interrelated,
that it is more important in framing genera than families or orders ;
more constant characters must be employed for the higher groups.
The general shape of the bill is referable to the cone; it is the
anterior part of the general cone that we have seen to reach from
its point to the base of the skull. This shape confers the greatest
strength combined with the greatest delicacy ; the end is fine to
apprehend the smallest objects, while the base is stout to manipulate
the largest. But in no bird is the cone expressed with entire pre-
cision ; and, in most, the departure from this figure is great. The
bill always consists of two, the upper and the lower

Mandibles (Fig. 26), which lie, as their names indicate, above
and below, and are separated by a horizontal fissure, — the mouth.
Each mandible always consists of certain
projecting skull-bones, sheathed with more
or less horny integument in lieu of true
skin. The framework of the Upper Man-
dible is (chiefly) a bone called the inter-
maxillary, or better, the premaxillary. In
general, this is a three-pronged or tripodal
bone running to a point in front, with
the uppermost prong, or foot, implanted

upon the forehead, and the other two, Fia.se.-Partsof aBm. «, side
lower and horizontal, running into the of upper mandible ; &, culmen ; c,

• 1 n . 1 -1 11 rpi PC 11 i! J.1 nssSii foss3/ \ nostril , c (s6g 1d6- .

sides 01 the skull. Ihe scanold 01 the low); /, gape, or whole commis-

Under Mandible is a compound bone

called inferior maxillary; it is U- or V- ^ ramus of under jaw; j, tomia of
shaped, with the point or convexity in lines e should have been drawn to
front and the prongs running to either S'*l“ppL‘mandibie)’rj^^^^^
side of the base of the skull behind, to
be there movably hinged. These two

bones, with certain accessory bones of the upper mandible, as the palate
bones, etc., together with the horny investment, constitute the Jaws.
Both jaws, in birds, are movable j the under, by the joint just men-
tioned ; the upper, either by a joint at, or by the elasticity of the
bones of, the forehead ; it is moved by a singular muscular and bony
apparatus in the palate, further notice of which is given beyond,
under head of Anatomy (Osteology). The motion of the upper
mandible is freest and most extensive in the parrot tribe, where
both fronto-maxillary and palato-maxillary sutures exist. \Yhen
closed, the jaws meet and fit along their apposed edges or surfaces.

GENERAL ORNITHOLOGY

PART II

in the same manner and for the same purposes as the lips and teeth
of man or other vertebrate animals. All bills, thus similarly consti-
tuted, have been divided by the author into

Four Classes, representing as many ways in which the two
mandibles close upon each other at the end. 1. The epignathous
(Gr. eiTL, epi, upon, yvdOos, gnathos, jaw) way, plan, or type, in which
the upper mandible is longer than the under, and its tij) is evidently
bent down over the tip of the lower. 2. The hypognathous (Gr.
P7TO, liupo, under), in which the lower mandible is longer than the
other. 3. The paragnathous (Gr. Tvapd, pam, at or by), in which
both are of about equal length, and neither is evidently bent over
the other. 4. The metagnathous (Gr. gerd, meta, with, beside, etc.),
in which the points of the mandibles cross each other. The second
and fourth of these are extremely rare ; they are exemplified, re-
spectively, by the skimmer and the crossbill (genera Ehynchops and
Loxia). The first is common, occurring throughout the birds of
prey, the parrots, and among the petrels, gulls, etc. etc. The great
majority of birds exhibit the third ; and, among them, there is such
evident gradation into epignathism, that it is necessary to restrict
the latter to its complete development, exhibited in the intermax-
illary bone divested of its horny sheath, which often, as among
flycatchers, etc., forms a little overhanging point, but does not
constitute epignathism. These classes, it should be added, though
always applicable, and very convenient in descriptions, are purely
arbitrary, that is, they by no means correspond to any four large
groups of birds ; but, on the contrary, usually only mark families
and the subdivisions of families ; and the four types may be seen
in closely related genera. The general shape of the bill has also
furnished

Other Classes, for many years used as a large basis for orni-
thological classification, even for the establishment of orders ; but
which the progress of the science has shown to be merely as con-
venient as, and only less arbitrary than, the foregoing. The principal
of these are represented by the following types : A, among land
birds. 1. The fissirostral (Lat. fissus, cleft, and rostrum), or cleft,
in which the bill is small, short, and with a very large gape running
down the side of the head ; as in the swallow, chimney-swift, goat-
sucker. ■ 2. The tenuirostral (Lat. tenuis, slender), or slender, in
which the bill is narrow, long, and with a short cleft; as in the
humming-bird, creeper, nuthatch. 3. The dentirostral (LaL dens, a
tooth), or toothed, in which, with a various general shape, there is
present a nick, tooth, or evident lobe in the apposed edges of one
or both mandibles near the end ; as in the shrike and some wrens,
thrushes, and warblers. 4. The conirostral (Lat. conus, a cone), or
conical, sufficiently defined by its name, and illustrated by the great

SEC. Ill

EXTERIOR FARTS OF BIRDS

149

finch or bunting family and some allied ones. — B, among water
birds. 5. The longirostral (Lat. longus, long), or long, an aquatic
style of the tenuirostral, best exhibited in the great snipe family.
6. The pressirostral (Lat. pressus, pressed), or the compact, illustrated
by the plovers, etc. 7. The cuUrirostral (Lat. culter, a knife), cut-
ting, exemplified in the heron group. 8. The lamellirostml (Lat.
lamella, a plate), in which the bill is broad and plated with a series
of tooth-like processes, as in the duck tribe. None of these terms
are now used to indicate natural groups, nor have we such absurd-
ities as the “ orders ” Fissirostres, Tenuirostres, etc. A swallow, for
instance, and a swift are equally fissirostral, though only distantly
related to each other ; a swift is closely related to a humming-bird,
though the latter is extremely tenuirostral ; and birds of contiguous
genera may be dentirostral or not. The words are nevertheless
convenient incidental terms in general descriptions. Various other
similar terms, expressing special modifications, as latirostral (Lat.
latus, broad), acutirostral (Lat. acutus, sharp), etc,, are also employed
as common adjectives.

Other Forms. — A bill is called long, when notably longer than
the head proper ; short, when notably shorter ; medium, in neither
of these conditions. It is compressed, when higher than wide, at
the base at least, and generally for some portion of its length ;
depressed, when wider than high ; terete (Lat. teres, cylindric), under
neither of these conditions. It is recurved, when curved upward j
decurved, when curved downward ; lent, when the variation in either
direction is at an angle ; straight, when not out of line with the
axis of the head. A bill is oUuse (said chiefly of the paragnathous
sort) when it rapidly comes to an end that therefore is not fine ; or
when the end is knobby ; it is acute when it runs to a sharp point ;
acuininate, when equally sharp and slenderer; attenuate, when still
slenderer ; subulate (awl-shaped), when slenderer still ; acicular
(needle-shaped), when slenderest possible, as in some humming-birds.
A bill is arched, vaulted, turgid, tumid, inflated, etc., when its outlines,
both crosswise and lengthwise, are notably more or less convex ;
and contracted, when some, or the principal, outlines are concave
(said chiefly of depressions about the base of the upper mandible,
or of concavity along the sides of both mandibles). A bill is
hamulate (Lat. hamus, a hook), or unguiculate (Lat. unguis, a claw),
when strongly epignathous, as in rapacious birds, where the upper
mandible is like the talon of a carnivorous beast ; it is dentate,
when toothed, as in a falcon ; if there are a number of similar
“ teeth,” it is serrate (Lat. serra, a saw), like a saw ; it is cultrate
(knife-like), when extremely compressed and sharp-edged, as in the
auk, skimmer : if much curved as well as cultrate, it is falcate (Lat.
falx, a reaping-hook ; scythe-shaped) ; and each mandible may be

GENERAL ORNITHOLOGY

PART II

150

oppositely falcate, as in the crossbill, constituting metagnathism.
A bill much flattened and widened at the end (rare) is spatulate
(Lat. spatula,, a spoon) ; examples : spoonbill, shoveller duck. One
is called lamellate, when it has a series of plates or processes just
inside the edges of the mandibles ; as in all the duck order, and in
a few petrels ; the design is to furnish a sifter or strainer of water,
just what is effected in the whale, by the baleen in its mouth.
Finally, the far end of the bill, of whatever shape, is called the tip
or ayex (Fig. 26, n) ; the near end, joined to the rest of the skull,
the base ; the rest is the continuity. Some other features of the bill
as a whole are best treated under the separate head of

The Covering of the Bill. — (a.) In the great majority of birds,
including nearly all perchers, many walkers, and some waders and
swimmers, the sheathing of the mandibles is wholly hard, horny, or
corneous (Lat. cornu, a horn) ; it is integument modified much as in
the case of the nails or claws of beasts. In nearly all waders, and
most swimmers, the sheath becomes, wholly or partly, softer, and
is of a dense, leathery texture. But some swimmers, as among the
auks, furnish bills as hard-covered as any, while some perchers have
it partly quite soft, so that no unexceptional rule can be laid down ;
and, moreover, the gradations from one extreme to the other are
insensible. Probably the softest bill is found among the snipes,
where it is skinny throughout, and in typical snipes and woodcocks
vascular and nervous at the tip, becoming a true organ of touch,
used to feel for worms out of sight in the mud. In all the duck
order the bill is likewise soft ; but there it is always terminated by
a hard, horny unguis or “ nail,” more or less distinct j and such a
horny claw also occurs in other water birds with softish bills, as the
pelican. An interesting modification occurs in all, or nearly all, of
the pigeon order ; these birds have the bill hard or hardish at tip
and through most of continuity, but towards and at the base of the
upper mandible the sheath changes to a soft, tumid, skinny texture,
overarching the nostrils ; it is much the same with most plovers.
But the most important feature in this connection is afforded by
the parrots and all the birds of prey : one so remarkable that it
has received a distinct name : Cere. The cere (Lat. cera, wax ;
because it looks waxy) is a dense membrane saddled on the upper
mandible at base, so different from the rest of the bill, that it might
be questioned whether it does not more properly belong to the head
than to the bill, were it not for the fact that the nostrils open in it.
Moreover, the cere is often densely feathered, as in the Carolina
paroquet, in the bill proper of which no nostrils are seen, these
being hidden in the feathered cere, which, therefore, might easily
be mistaken at first sight for the bird’s forehead. A sort of false
cere occurs in some water birds, as the jaegers, or skua-gulls (genus

SEC. Ill

EXTERIOR FARTS OF BIRDS

151

Stercorarius). The tumid nasal skin of pigeons is sometimes called
a cere ; but the term had better be restricted to the birds first
above named. The under mandible probably never presents soften-
ing, except as a part of general skinniness of the bill ; it may have
a nail at the end. (b.) The covering is either entire or pieced. In
most birds it is entire ; that is, the sheath of either mandible may
be pulled off whole, like the finger of a glove. It is, however, in
many birds divided into parts, by various lines of slight connection,
and then comes off in pieces ; as is the case with some water birds,
particularly petrels, where the divisions are regular, and the pieces
have received distinctive names. Many auks {Alcidce) have the
covering of the bill in particular pieces, and it is an extra-
ordinary fact that such parts are of a secondary sexual character,
being assumed at the breeding season and afterwards moulted like
feathers. Such condition of the sheath of the beak, or of special
developments of the sheath, is called caducous or deciduous. The
entire covering of both jaws together is called rliamphotlieca (Gr.
pdfjicfios, hramphos, beak ; theke, a sheath) ; of the upper alone,

rhmotheca (Gr. pis, hris, the nose) ; of the under, gnathotheca (Gr.
yvdOos, gnccthos, jaw) ; but these terms are not much used, (c.)
The covering is otherwise variously marked ; sometimes so strongly
that similar features are impressed upon the bones themselves be-
neath. The most frequent marks are various ridges (Lat. pi. carince,
keels) of all lengths and degrees of expression, straight or curved,
vertical, oblique, horizontal, lengthwise, or transverse; a bill so
marked is said to be striate (Lat. stria, a streak) or carinate ; when
numerous and irregular, they are called rugce (Lat. ruga, a wrinkle)
and the bill is said to be corrugated or rugose. When the elevations
are in points or spots instead of lines, they are called puncta (Lat.
punctum, a point) ; a bill so furnished is punctate, but the last word
is oftener employed to designate the presence of little pits or de-
pressions, as in the dried bill of a snipe towards the end. Larger
softish, irregular knobs or elevations pass under the general name
of warts or 'papillce, and a bill so marked is papillose ; when the
processes are very large and soft, the bill is said to be carunculate
(Lat. caro, flesh, diminutive carunculus, little bit of flesh). Various
linear depressions, often but not always associated with carinse, are
grooves or sulci (Lat. sulcus, a furrow), and the bill is then called
silicate. Sulci, like carinse, are of all shapes, sizes, and positions ;
when very large and definite, they are sometimes called canaliculi,
or channels. The various knobs, “ horns,” and large special features
of the bill cannot be here particularised. Any of the foregoing
features may occur on both mandibles, and they are exclusive of
that special mark of the upper (the nasal fossa) in which the
nostrils open, and which is considered below. We have still to

52

GENERAL ORNITHOLOGY

PART II

notice the special parts of either mandible j and will begin with
the simplest, the

Under Mandible. — In the majority of birds it is a little shorter
and a little narrower and not nearly so deep as the upper ; but
sometimes quite as large, or even larger. The upper edge, double
{i.e. there is an edge on both sides), is called the mandibular tomium,
or in the plural, tomia (Gr. refxveLv, temnein, to cut ; Fig. 26, j), as
far as it is hard ; this is received against, and usually a little within,
the corresponding edge of the upper mandible. The prongs already
mentioned are the mandibular rami (pi. of Lat. ramus, a branch ;
Fig. 26, i) these meet at some point in front, either at a short
angle (like >) or with a rounded joining (like ID). At their point of
union there is a prominence, more or less marked (Fig. 26, k) , this
is the gonys. That is to say, this point is gonys proper : but the
term is extended to apply to the whole line of union of the rami,
from gonys proper to the tip of the under mandible j and in de-
scriptions it means, then, the under outline of the hill for a corre-
sponding distance (Fig. 26, 1). This important term must be
understood ; it is constantly used in describing birds. The gonys
is to the under mandible what the keel is to a boat ; it is the
opposite of the ridge or culmen of the upper mandible. It varies
greatly in length. Ordinarily it forms, say, one -half to three-
fourths of the under outline. Sometimes, as in conirostral birds,
a sparrow, for example, it represents nearly all this outline ; while
in a few birds it makes the whole, and in some, as the puffin, is
actually longer than the lower mandible proper, because it extends
backwards in a point. Other birds may have almost no gonys at
all j as a pelican, where the rami only meet at the extreme tip, or
in the whole duck family, where there is hardly more. As the
student must see, the length of the gonys is simply a matter of
how extensive is the fusion of the rami ; and that, similarly, their
mode of fusion, as in a sharp ridge, a flat surface, a straight line,
a curve, etc., results in corresponding modiflcations of its special
shape. The interramal space is complementary to length of gonys :
sometimes it runs to the tip of the bill, as in a pelican, sometimes
there is next. to none, as in a puffin; while its width depends upon
the degree of divergence, and the straightness or curvature, of the
rami. The surface between the tomium and lower edge of rami
and gonys together is the side of the under mandible (Fig. 26, m).
The most important feature of the

Upper Mandible is the culmen (Lat. for top of anything ; Fig.
26, h). The culmen is to the upper mandible what the ridge is to
the roof of a house ; it is the upper profile of the bill — the highest
middle lengthwise line of the hill ; it begins where the feathers end
on the forehead, and extends to the tip of the upper mandible.

SEC. Ill

EXTERIOR TARTS OF BIRDS

153

According to the shape of the bill it may be straight or convex, or
concave, or even somewhat m -shaped ; or double-convex, as in the
tufted puffin : but in the great majority of cases it is convex, with
increasing convexity towards the tip. Sometimes it rises up into
a thin elevated crest, as well shown in the genus Crotoyhaga, and
in the puffins {Fmtercula), when the upper mandible is said to be
keeled, and the culmen itself to be cultmte ; sometimes it is really a
furrow instead of a ridge, as toward the end of a snipe’s bill ; but
generally it is simply the uppermost line of union of the gently
convex and sloping sides of the upper mandible (Fig. 26, a). In a
great many birds, especially those with depressed bill, as all the
ducks, there is really no culmen j but then the median lengthwise
line of the surface of the upper mandible takes the place and name
of culmen. The culmen generally stops short about opposite the
proper base of the bill ; then the feathers sweep across its end, and
downwards across the base of the sides of the upper mandible,
usually also obliquely backwards. Variations in both directions
from this standard are frequent ; the feathers may run out in a
point on the culmen, shortening the latter, or the culmen may run
a way up the forehead, parting the feathers ; either in a point, as
in the rails and gallinaceous birds, or as a broad plate of horn, as
in the coots and gallinules. The lower edge (double) of the upper
mandible is the maxillary tomium, as far backward as it is hard and
horny. The most conspicuous feature of the upper mandible in
most birds is the

Nasal Fossa (Lat. fossa, a ditch), or nasal groove (Fig. 26, c), in
which the nostrils open. The upper prong of the intermaxillary
bone is usually separated some ways from the two lateral prongs ;
the skinny or horny sheath that stretches betwixt them is usually
sunken below the general level of the bill, especially in those birds
where the prongs are long or widely separated ; this “ ditch ” is
what we are about. It is called fossa when short and wide, with
varying depth ; sulcus or groove when long and narrow ; the
former is well illustrated in the gallinaceous birds ; the latter in
nearly all wading birds and many swimmers. AVhen the inter-
maxillary prongs are soldered throughout, or are very short and
close together, there is no (or no evident) nasal depression, the
nostrils then opening flush with the level of the bill. The

Nostrils (Fig. 26, d), two in number, vary m ^position as follows :
— they are lateral, when on the sides of the upper mandible
(almost always) j culminal, when together on the ridge (rare) ;
superior or inferior when evidently above or below midway betwixt
culmen and tomia ; they are lasal, when at the base of the upper
mandible ; suh-hasal when near it (usual) ; median when at or near
the middle of the upper mandible (frequent, as in cranes, geese.

154

GEJVERAL ORNITHOLOGY

PART II

etc.); terminal when beyond this (very rare; probably there are
now no birds with nostrils at the end of the bill, except the
Apteryx). The nostrils are pervious, when open, as in nearly all
birds ; impervious, when not visibly open, as among cormorants and
other birds of the same order ; they are perforate when there is no
septum (partition) between them, so that you can look through them
from one side of the bill to the other, as in the turkey-buzzard,
crane, etc.; imperforate when partitioned off from each other, as in
most birds ; but different ornithologists use these terms interchange-
ably. The principal shapes of the nostrils may be thus exhibited : —
a line, linear nostrils ; a line variously enlarged at either end,
clavate, club-shaped, oblong, ovate nostrils; a line, enlarged in the
middle, oval or elliptic nostrils ; this passing insensibly into the
circle, round or circular nostrils ; and the various kinds of more or
less linear nostrils may be either longitudinal, as in most birds, or
oblique, as in a few; very seldom directly transverse (up and down).
Rounded nostrils may have a raised border or rim; when this is
prolonged they are called tubular, as in some of the goatsucker
family, and in all the petrels. Usually, the nostrils are defined
entirely by the substance surrounding them ; thus, of cere, in a
hawk ; of softish skin, in a pigeon, plover, or snipe ; or of horn, in
most birds ; but often their contour is partly formed by a special
development, somewhat distinct either in form or texture, and this
is called the nasal scale. Generally, it forms a sort of overhanging
arch or portico, as well shown in all the gallinaceous birds, among
the wrens, etc. A very curious case of this is seen in the wryneck
{lynx torquilla), where the scale forms the floor instead of the roof
of the nostrils. The nostrils also vary in feathered or naked ;
the nasal fossa being a place where the frontal feathers are apt to
run out in points (called antice), embracing the root of the culmen.
This extension may completely fill and hide the fossa, as in many
grouse and ptarmigan ; but it oftener runs for a varying distance
toward, or above and beyond, the nostrils ; sometimes similarly
below them, as in a chimney-swift ; and the nostrils may be densely
feathered when there is no evident fossa, as in an auk. When thus
truly feathered in varying degree, they are still open to view ;
another condition is, their being covered over and hidden by
modified feathers not growing on the bill itself, but on the forehead.
These are usually bristle-like {setaceous), and form two tufts, close-
pressed and directed forwards, as is perfectly shown in a crow ; or,
the feathers may be less modified in texture, and form either two
tufts, one over each nostril, or a single ruff, embracing the whole
base of the upper mandible ; as in nuthatches, titmice, red-poll
linnets, snow -buntings, and many other northern Fringillidm.
Bristles or feathers thus growing forward are called retrorse (Lat.

SEC. Ill

EXTERIOR TARTS OF BIRDS

155

retrorsum, backward; here used in the sense of in an op;posite
direction from the lay of the general plumage ; but they should
properly be called antrorse, i.e. forward). The nostrils, whether
culminal or lateral, are, like the eyes and ears, always two in
number, though they may be united in one tube, as in the petrels.

The Gape. — It only remains to consider what results from the
relations of the two mandibles to each other. When the bill is
opened, there is a cleft or fissure between them ; this is the ga'pe or
rictus (Lat. rictus^ mouth in the act of grinning). But while thus
really meaning the open space between the mandibles, it is generally
used to signify the line of their closure. Commissure (Lat. committere, to
put or join together) means the point where the gape ends behind,
that is, the angle of the mouth, angulus oris, where the apposed
edges of the mandibles join each other ; but, as in the last case, it
is loosely applied to the whole line of closure, from true commissure
to tip of the bill. So we, say, “commissure straight,” or “commis-
sure curved ” ; also, “ commissural edge ” of either mandible
(equivalent to “ tomial edge ”) in distinction from culmen or gonys.
But it would be well to have more precision in this matter. Let,
then, tomia (Fig. 26, 7) be the true cutting edges of either mandible
from tip to opposite base of bill proper; rictus (Fig. 26, g) be their
edges thence to the point commissure (Fig. 26, h) where they join
when the bill is open; the line commissure (Fig. 26, f) to include
both when the bill is closed. The gape is straight, when rictus and
tomia are both straight and lie in the same line ; curved, sinuate,
when they lie in the same curved or waved line ; angulated, when
they are straight, or nearly so, but do not lie in the same line, and
therefore meet at an angle. Angulation of the commissure is a
distinctive character of most finches and buntings — Fringillidce. It
is well shown in the hawfinch, Coccothraustes mdgaris.

II. THE WINGS.

Definition. — Pair of anterior or pectorcd limbs organised for
flight by means of dermal outgrowths. Used for this purpose by
birds in general ; but by ostriches and their allies only as outriggers
to aid running ; by penguins as fins for swimming under water ;
used also in the latter capacity by some birds that fly well, as
loons, cormorants, dippers. Wanting in no recent birds, but im-
perfect in a few, as all Ratitce ; greatly reduced in the emeu, casso-
wary, and kiwi-kiwi ; also in the moas {Dinornis) ; in the Cretaceous
Hesperornis only the rudimentary humerus is known. To under-
stand their structure we must notice particularly

The Bony Framework (Figs. 27, 28, 29). — The skeleton of a
bird’s wing is built upon a plan common to the fore or pectoral

GENERAL ORNITHOLOGY

PART II

156

limb of all the higher vertebrates, so that its bones and joints may
readily be compared and identified with those of any lizard or
mammal, including man. But this member is highly specialised ;

Fig. 27. — Bones of right wing of a duck, Clangula islandica, from above, | nat. size. (Dr.
R. W. Shufeldt, U.S.A.) A, shoulder, omos; R, elbow, ancon; C, wrist, carpus; D, end of
principal finger ; E, end of hand proper, metacarpus. A B, upper arm, hrachium ; B C, fore-
arm, antebrachium ; C D, whole hand or pinion, manus; composed of C E, hand proper or meta-
carpus, excepting d 2 ; F D, or d 2, d 3, d 4, fingers, digits, digiti. Ti, humerus ; rd, radius ; ul,
ulna ; sc, outer carpal, scapholunare or radiate ; cu, inner carpal, cuneiforms or ulnare ; these
two composing wrist or carpus, me, the compound hand-bone, or metacarpus, composed of three
metacarpal bones, bearing as many digits — the outer digit seated upon a protuberance at the
head of the metacarpal, the other two situated at the end of the bone, d 2, the outer or radial
digit, commonly called the thumb or pollex, composed of two phalanges ; d 3, the middle digit,
of two phalanges ; d the inner or ulnar digit, of one phalanx ; d 2 is the seat of the feathers of
the bastard wing or alula. D to C (whole pinion), seat of the fiight-feathers called primaries ; C
to B (forearm), seat of the secondaries; at B and above it in direction of A, seat of tertiaries
proper ; below A, in direction of B, seat of scapularies (upon pteryla humeralis), often called
tertiaries. The wing shown half-spread : complete extension would bring ABC D into a right
line ; in complete folding C goes to A, and D to B; all these motions nearly in the plane of the
paper. The elbow-joint and wrist are such perfect hinges, that, in opening or closing the wing,
C cannot sink below the paper, nor D fly up above the paper, as would otherwise be the effect of
the pressure of the air upon the flight-feathers. Observe also : rd and ul are two rods con-
necting B and C ; the construction of their jointing at B and C, and with each other, is such
that they can slide lengthwise a little upon each other. Now when the point C, revolving about
B, approaches A in the arc of a circle, rd pushes on sc, while ul pulls back cu ; the motion is
transmitted to D, and makes this point approach B. Conversely, in opening the wing, rd pulls
back sc, and ul pushes on cu, making D recede from B. In other words, the angle ABC cannot
be increased or diminished without similarly increasing or diminishing the angle R CD; so
that no part of the wing can be opened or shut without automatically opening or shutting the
rest, — an interesting mechanism by which musciilar power is correlated and economised. This
latter mechanism is further illustrated in Fig. 28, where rc and uc show respectively the size,
shape, and position of the radial condyle and ulnar condyle of the humerus. It is evident that
in the flexed state of the elbow, as shown in the middle figure, the radius, rd, is so pushed upon
that its end projects beyond ul, the ulna ; while in the opposite condition of extension, shown
in the lower figure, rd is pulled back to a corresponding extent.

being fitted for accomplishing fiight, not only by the development
of feathers, but also by modifications in the bones themselves. The
axes of the bones have a special direction with reference to each

SEC. Ill

EXTERIOR PARTS OF BIRDS

157

other and to the axes of the body ; the movements of the joints
are peculiar in some respects ; and the whole extremity of the
wing, from the wrist outward, is peculiarly constructed, by loss of
some of the digits that five-fingered animals possess, and by the
compression of those that are left. The wing proper begins at the
shoulder- joint, where it hinges freely upon the shoulder, in a
shallow or glenoid socket formed conjointly by the shoulder-blade
or scapula, and by the coracoid bone ; these two, with the clavicles,
collar-bones, or merrythought, furculum, forming the shoulder-girdle,
or pectoral arch (Figs. 56, 59).

The wing ordinarily consists, in adult life, of ten or eleven
actually separate bones ; in the embryo (see Fig. 29) there are
indications of several more at the wrist-joint, which speedily lose

their individual identity by fusing together and with bones of the
hand.^v^Aside from these, there is often an accessory ossicle at the
shoulder-joint (Fig. 56, ohs), sometimes one at the wrist -joint,
occasionally an extra bone at the end of the principal finger. The
normal or usual number is shown in Fig. 27, taken from a duck
{Clangula islandica), in which there are eleven.

The upper-arm bone h, reaching from the shoulder A to the
elbow B, is the humerus. In the closed wing, the humerus lies
nearly in the position of the same bone in man when the elbow is
against the side of the body ; in extension of the wing the elbow
is borne away from the body, as when we raise the arm, but carry
it neither forward nor backward. A peculiarity of the bird’s
humerus is, that it is rotated on its axis through about the quadrant
of a circle, so that what is the front of the human bone is the outer

GENERAL ORNITHOLOGY

PART II

158

aspect in the bird. The humerus is a cylindric bone, straightish or
somewhat italic-/-shaped, with a globular head to fit the socket of
the shoulder, a strong pectoral ridge for insertion of the breast
muscles, and at the bottom two condyles (Fig, 28, rc, uc) or joint-
surfaces for articulation with a pair
of succeeding bones. The forearm,
cubit or anfebrachiwn, extending from
elbow to wrist, B to (7, in Fig. 27,
has two parallel bones of about
equal lengths. These are the ulna,
id, and the radius, rd ; the former,
inner and posterior, the larger of
the two, bearing the quills of the
secondary series ; the latter, slen-
derer, outer, and anterior. The
enlarged proximal extremity of the
ulna is called the olecranon, or,
“ head of the elbow.” The third
segment of the wing is the wrist or
carpus. In adult life, this normally
consists of two little knobby carpal
bones, extremely irregular in shape,
called the scapliolunare, sc, and cunei-
forme, cu. One being at the end of
the radius, the other at that of the
ulna, they are also called radiale

Fio.2S»,froma!,o«n5coek.o(.the.piains ^nd uhwre. In the embryo there
(Centrocercus urophasianus, six months old) ig ^t least One Other Carpal bone,
is designed to show the composition of the i r • i i

carpus and metacarpus before the elements that early lUSeS With the next
of these bones fuse together : r, radius ; u, , i i i j. i

ulna ; s, scapholunar or radiale ; c, cunei- Segment, and in many birds there

are several such. This fourth seg-

to be os magnum, later fusing with the ^ o

metacarpus ; 2;, a carpal bone, supposed to ment is the hand proper, or mcta-
be unciform, later fusing with metacarpus ; -n / ^ • o i

8, an unidentified fifth carpal bone, which CarpUS, lllC, C tO B (exclUSlVe 01 d

2). The single metacarpal or hand-

carpal bone bearing the pollex or outer j^QUe is Very Composite ; that is,
digit, consisting of two phalanges, d and A: ; ^ ’

9', principal O^iedian) metacarpal bone, compounded 01 Several ; tor, besides

wrpCang”ef!iyS?r9rrnSe?ol including certain carpal elements,
as already said, it consists of three

fuse with 9' to form the single compound bones fused (in all recent birds) in
metacarpal bone marked me in Fig. 2T. ^

(Prom nature, by Dr. R. w. shufeidt. One, Corresponding to the three
digits or fingers that birds possess.
In fact it is three metacarpals in one. The metacarpal corre-
sponding to the principal finger is much the largest of the
three ; that of the first finger is very short, being only the
expanded part seen in the figure, just above the bone marked

SEC. Ill

EXTERIOR FARTS OF BIRDS

159

d 2 j that of the third finger is nearly as long as the main
metacarpal, but much slenderer, and usually fused only at its two
ends, leaving between itself and the main metacarpal a considerable
space, as seen opposite the letters me in the figure. The wing is
finished off with three fingers or digits marked d 2, d d 4:. The
middle one of these, E to D m the figure, is much the largest, and
forms the main continuation of the hand. The digit, d 3, ordinarily
consists of two bones, cdX\&di phalanges, placed end to end, as in the
example before us ; but occasionally there is found a third phalanx.
The outer or radial digit, d 2, ordinarily consists of two bones, of

Fig. 29 Z?is.— Diagram of fore limbs of man, bat, horse, and bird. The lines 1-9 are isotomes,
cutting the limbs into morphologically equal parts, or isomeres.

which the terminal one is small, and may be wanting. The inner
or ulnar digit, d 4, consists of a single small phalanx, closely bound
to the side of the middle finger. Corresponding to the compactness
and consolidation of these terminal segments, the digits enjoy
little individual motion. The outer or radial digit is the most
independent one. In the Archceopteryx the three metacarpals were
free bones, and the whole hand more like that of a lizard. No bird
now has free metacarpals in adult life j none has more than three
digits. These three are supposed by some to correspond to the
thumb and fore and middle fingers of our hands ; by others, to the

i6o

GENERAL ORNITHOLOGY

PART II

fore, middle, and ring fingers, and being consequently the second,
third, and fourth digits, as marked in the figure. The digit marked
6^ 2 is commonly called a bird’s thumb or pollex. The Apteryx and
the cassowaries have but one complete digit. The resemblance to
a lizard’s or quadruped’s digits is increased by the claws which many
birds possess, These may be borne on the enlarged terminal
phalanx oi d 2 (Jc, in Fig. 29), as is very well shown in the turkey-
buzzard and other American Cathartidce ; both on this and on the
terminal phalanx of c? 3 {d" in Fig. 29), as in the ostrich ; on the
latter alone, as in the Apteryx, cassowary, American ostrich, and
swan. The inner finger, d 4 {d'" in Fig. 29) is not known to ever
bear a claw, excepting in Archceopteryx. The whole segment, C to
D, is commonly called “ the hand,” “ pinion,” or manus, though, as
we have seen, it consists of hand proper (metacarpus), and fingers
(digits) with their respective phalanges. (Fig. 29 bis.)

Some other bones are observed in birds’ wings. As already
said, there is a supplementary ossicle in the shoulder-joint of many
birds ; it is badly called the scapula accessoria (Fig. 56, ohs). At the
convexity of the elbow there may be one or more ossicles, not per-
taining properly to the wing-skeleton, but developed in the tendons
of muscles passing over the joint ; they are sesamoids, like the
human patella, or knee-cap. In various birds there is found at
the convexity of the wrist, on the head of the metacarpal, an
ossicle called the osprominens ; apparently a sesamoid. Some other
ossicles observed in the wrists of young or embryonic birds are all
supposed to be carpal elements, the exact homologies of which may
be still questioned.

The Mechanism of these Bones is admirable. The shoulder-
joint is free, much like our own, permitting the humerus to swing
all about ; though the principal motions are to and from the side of
the body {adduction and abduction), and up and down in a vertical
plane. The elbow-joint is a very strict hinge, permitting motion in
one plane, nearly that of the wing itself. The finger-bones have
little individual motion. The construction of the wrist-joint is
quite peculiar. In the first place the two bones of the forearm are
so fixed in relation to each other, that the radius cannot roll over
the ulna, like ours. If you stretch your arm upon the table, you
can, without moving the elbow, turn the hand over so that either
the palm or the knuckles are downward. This is a rotary motion
of the bones of the forearm, called pronation and supination; the
hand is prone when the palm touches the table, supine when the
knuckles are downward. This rotation is absent from the bird’s
arm ; if it could occur, the action of the air upon the pinion-
feathers would throw them all “ at sea ” during the strokes of the
wing, rendering flight difficult or impossible. The hinging of the

SEC. Ill

EXTERIOR PARTS OF BIRDS

i6i

hand upon the wrist is such, also, that the hand does not move up
and down, as ours can, in a plane perpendicular to the surface of
the wing, but in the same plane as that surface. The motion is
that which would take place in our hand if we could bring the little
finger and its border of the hand so far around as to touch the
corresponding border of the forearm. It is a motion of adduction,
not of flexion, and its opposite, abduction, not extension, by which
a wing is folded and spread. Such abduction is the way in which
the hand is extended upon the wrist -joint, increasing and com-
pleting the unfolding of the wing that begins by the true extension
of the forearm upon the elbow and abduction of the upper arm from
the body. In a word, a wing is spread by the motion of abduction
at the shoulder and wrist, of extension at the elbow j it is closed
by adduction at the shoulder and wrist, and flexion at the elbow.
The numerous muscles which unfold or straighten out the wing are
called extensors ; those that bend or close it are flexors. Extensors
lie upon the back of the upper arm, and the front of the forearm
and hand, their “ leaders ” or tendons passing over the convexities of
the elbow and of the wrist. The flexors occupy the opposite sides
of the limb, with tendons in the concavities of the joints. The
most powerful muscles of the wings are the great pectoral or breast
muscles, acting upon the upper end of the humerus j there are
several of them, exerted in throwing out the arm from the body,
and in giving both the up and down wing-strokes. Tendons are
generally strong inelastic cords ; but there is an interesting arrange-
ment of an elastic cord in a bird’s wing. In Fig. 27, A B C is a
deep angle formed by the naked bones, but none such is visible from
the exterior, because the space is filled by a fold of skin passing
from G to near A. But C approaches and recedes from A as the
wing is folded or unfolded, and a cord long enough to reach A-C
would be slack in the folded wing, did not its elasticity enable it to
contract and stretch, keeping the anterior border of the wing straight
and smooth. (For another automatic mechanism, see Fig. 28.)

The point (7 is a highly important landmark in practical orni-
thology it represents, in any folded wing, a very prominent point,
the distance from which to the tip of the longest flight-feather is a
special measurement known as that of “the wing.” It is the con-
vexity of the carpus, commonly called the “ carpal angle,” or “ bend
of the wing.” Having thus glanced at. the bony structure and
mechanism of the wing, we are ready to examine the

Feathers of the Wing* (Fig. 30). — How important these are will
be evident from the consideration that they are the bird’s chief
organs of locomotion ; for without them the wing would be useless
for flight. We also remember that such means of locomotion is the
great specialty of birds. Wing-feathers are those which grow upon
M

GENERAL ORNITHOLOGY

PART II

162

the ^teryla alaris. They are of two main sorts : the flight-feathers
proper, or long stiff quills, collectively called remiges (Lat. remex, pi.
remiges, rowers) ; and the smaller, weaker feathers overlying them,
and hence called coverts, or tectrices (Lat. tectrix, pi. tectrices, coverers).
To these may be added as a third distinct group, the bastard quills,
which constitute the

Alula, OP Ala Spupia (Lat. alula, little wing, diminutive of ala,
wing ; spuria, spurious, bastard). The “ little wing ” is simply the
small parcel of feathers which grow upon the “ thumb ” (see Fig. 27,
d 2 2^, d and k ; 30, al). Highly significant as these may be in

a morphological point of view, as representing what this part of the
wing may have been in early times, they are so much reduced in
modern birds as to be of little account in practical ornithology. In
fact^ the unpractised student may fail to recognise them at first.
They form a small packet on the fore outer border of the pinion
near the carpal angle, and lie smoothly upon the upper surface of
the wing, strengthening and finishing off what would be otherwise
a weak spot in the contour of the wing-border. It is quite easy, on
recognising them, to lift them collectively a little aw^ay from the
other feathers, owing to the mobility of thumb. In fact, they are
sometimes quite obtrusive, when faulty taxidermy has discomposed
them. They are not often conspicuously modified either in size or
colour. In a few birds {e.g. Cathartes), a claw will be found at the
end of the joint which bears them. (The student must be careful
to discriminate between the use of the word spurious in the present
connection and its application to a rudimentary condition of the
first remex ; see p. 167.) The

Wing-Covepts overlie the bases of the large quills on both
the upper and under surfaces of the wing. They are therefore
conveniently divided into an upper set {tectrices superiores) and an
under set {ted. inferiores). The former are so much more conspicuous
than the latter that they are always understood when “ upper ” is
not specified. The latter are sometimes collectively called “ the
lining of the wings.” Coverts include all the small feathers of the
wings excepting the bastard quills ; they extend a varying distance
along the bases of the flight-feathers. The ordinary disposition
and division of the upper coverts is as follows : — One set, rather
long and stiffish, grow upon the pinion, and are close-pressed upon
the bases of the outer nine or ten remiges, covering these large
feathers about as far as their structure is plumulaceous. These are
the upper primary coverts, or coverts of the primaries (Fig 30, pc) ;
they are ordinarily the least conspicuous of any. All the rest of
the upper coverts are secondary ; they spring mostly from the
forearm. These are considered in three groups or rows. The
greater upper secondary coverts, called simply the “ greater coverts ”

SEC. Ill

EXTERIOR FARTS OF BIRDS

163

(tectrices majores, Fig. 30, gsc), are the first, outermost, longest row,
reaching nearest the tips of the flight-feathers ; they overlie the
bases of nearly all the remiges, excepting the first nine or ten. The
median upper secondary coverts, shortly known as the “ middle coverts ”
{tectrices medice), are a next row, shorter and therefore less exposed,
but still quite evidently forming a special series (Fig. 30, msc). It is a
common feature of these median coverts that they shingle over each
other contrariwise to the way the greater coverts are imbricated, the
outer vane of one being under the inner vane of the next outer one.
All the rest of the upper secondary coverts, forming several indis-
tinguishable rows, pass under the general name of lesser coverts
{tectrices minor es ; Fig. 30, he). The greater coverts furnish an excel-

Fig. 30. — Feathers of a sparrow’s wing ; nat. size. (For explanation see text.)

lent zoological character ; for in no Passeres are they more than half
as long as the remiges they cover, while the reverse is the case in
most birds of lower orders. Woodpeckers, however, though non-
passerine, have quite short coverts. The under coverts have the
same general arrangement as the upper ; but they are more alike
and less distinctly disposed in rows or series ; so that for practical
purposes they pass under the general name of under wing-coverts, or
lining of the wing. Since, when the wing is particularly marked on
the under side, it is the coverts and not the remiges that are highly
or variously coloured, the common expression “ wing below,” or
“under surface of the wing,” refers to these coverts more particu-
larly. We should distinguish, however, from the under coverts in

164

GENERAL ORNITHOLOGY

PART II

general, the axillars, or axillary feathers (Lat. axilla, the armpit).
These are the innermost feathers lining the wings, lying close to the
body; almost always longer, stiffer, narrower, or otherwise pecu-
liarly modified. In ducks, for example, and many of the waders, as
snipe and plover, they are remarkably well developed. The colour
of the axillaries is the principal distinction between some species of
plovers. The

Remiges, or Flight-Feathers (Fig. 30, h, s, and t), give the wing
its general character, mainly determining both its size and its shape ;
they represent most of its surface and of its inner and outer borders,
and all of its posterior outline, forming a great expansion, of which the
bony and fleshy framework is insignificant in comparison. The
shape of the wing is indeed primarily affected by the relative
lengths of its bony segments, the upper arm being, in a humming-
bird, for example, very short in comparison with the terminal
portion of the limb, and in an albatross again, both upper and fore-
arm being greatly lengthened ; still in any case it is the flight-
feathers that mainly determine the contour of the wing, by their
absolute degree of development, their lengths proportionately to
one another, and their individual shapes. They collectively form a
thin, elastic, flattened surface for striking the air, quite firm along
the front border where the bone and muscle lie, thence growing
more mobile and resilient toward the posterior border and along
the outer edge. Such surface may be quite flat, as in such birds as
cut the air with long, pointed wings, like oar-blades ; but it is
generally a little concave underneath, and correspondingly convex
above ; such arching or vaulting of the wing-surface being usually
associated with a short, broad, rounded wing, as in the gallinaceous
tribe, and being least in birds which have the thinnest and sharpest
’^Wngs. Corresponding differences in the mode of flight result.
The short, rounded wing confers a powerful though laboured flight
for short distances, usually accompanied by a whirring noise result-
ing from the rapidity of the wing-beats ; birds that fly thus are
almost always thickset and heavy. The long, pointed wing gives a
noiseless, airy, skimming flight, indefinitely prolonged, and accom-
plished with more deliberate wing-beats ; birds of this style of
wing are generally trim and elegant. These, of course, are merely
generalisations of the extremes of modes of flight, mixed and
gradated in every degree in actual bird-life. Thus the humming-
bird, which has sharp, thin wings, whirs them fastest of all birds, —
so rapidly that the eye cannot follow the strokes, merely perceiving
a haze about the bird while the ear hears the buzzing. The com-
bination of acuteness and concavo-convexity is a remarkably strong
one, conferring a rapid, vigorous, whistling flight, as that of a duck
or pigeon, or the splendid hurtling of a falcon. An ample wing, as

SEC. Ill

EXTERIOR TARTS OF BIRDS

165

one both long and broad without being pointed is called, is well
displayed by such birds as herons, ibises, and cranes ; the flight may
be strong and sustained, but is rather slow and heavy. The longest-
winged birds are found among the swimmers, particularly the
pelagic family of the petrels, and some of the whole-webbed order,
as pelicans, particularly the frigate-pelican. The last-named, Tadiy-
^etes aquilus, has perhaps the longest wings for its bulk of body of
any bird whatever, as well as the shortest feet. The American
vultures are likewise of great alar expanse in proportion to their
weight. The shortest wings, among birds possessing perfect remiges,
occur among the lower swimmers, as auks and divers, and among
some of the Gallince. The great auk is, or was, perhaps the only
flightless bird with well-formed flight-feathers, only too small to
subserve their usual purpose ; though certain South American ducks
are said to be in similar predicament. In the penguins, the whole
wing -structure is degraded, and the remiges abort in scale -like
feathers, the wings being reduced to fins both in form and function.
The whole of the existing Batitce or struthious birds, as the ostrich,
cassowary, and emeu, have rudimentary or very imperfect wings, as
was the case with the Cretaceous Hesperornis ; but the contem-
porary of the latter, Ichthyornis, and the still more ancient Archoeo-
^teryx, appear both to have had excellent ones.

The disposition of the remiges in their mutual relations is very
noteworthy. They have a rigid hollow barrel of great resistant
powers, considering the amount of substance, — just like the cylin-
drical stem of the cereal plant ; a stout, solid, highly-elastic shaft ;
the outer web narrower than the inner, with its barbs set at a more
acute angle upon the shaft. Any one of these stiffer outer vanes
overlies the broader and more yielding inner vane of the next outer
feather, which, on receiving the impact of air from below, resists as
it were with the strength of a second shaft superimposed. Though
the “ way of an eagle in the air ” was a mystery to the wise man of
old, the mechanics of ordinary flight are now better understood.
But the sailing of some birds for an indefinite length of time, up as
well as down, without visible motion of the wings, and without
reference to the wind, remains an enigma. The flight of the alba-
tross and turkey vulture, I venture to affirm, is not yet explained.
The riddle of The Wing will be read when we know how the arch-
saurian escaped from ilus to aether.

The number of true remiges ranges from about sixteen, as in a
humming-bird, to upwards of fifty, as in the albatross. Their shape
is quite uniform, minor details aside. They are the stiffest,
strongest, most perfectly fennaceous of feathers, without evident
hyporhachis, if any. They are generally lanceolate, that is, tapering
regularly and gradually to an obtuse point, though not infrequently

GENERAL ORNITHOLOGY

PART II

1 66

more parallel-sided, especially those of the secondary and tertiary
series. Either or both webs may be incised toward the end ; that
is, more or less abruptly narrowed ; this is called emargination ;
their ends may be transversely or obliquely truncate, or nicked
in various ways. In a few birds, apparently for purposes of
sexual ornamentation, they are developed in bizarre shapes of
beauty, with evident decrease of utility as flight-feathers. Those
of the ostrich and penguin tribes share the peculiarities of the
general plumage of these extraordinary birds. Eemiges are divided
into three classes or series, according to where they grow upon the
limb, whether upon the hand, the forearm, or the upper arm. In
this distinction is involved one of the most important considerations
of practical ornithology, of which the student must make himself
master. The three classes of quill-feathers are: th.Q jgnmaries ;

2, the secondaries ; 3, the tertiaries.

The Primaries (Fig. 30, b) are those remiges which grow upon
the pinion, or wrist-, hand-, and finger-bones collectively (Fig. 27, C
to D). Whatever the total number of the remiges may be, in nearly
all birds with true remiges the Primaries are either nine or ten in
number. The humming-bird with sixteen remiges, the albatross
with fifty or more, each has ten primaries. The grebes and a
few other birds are said to have eleven primaries : if this be so, it
is highly exceptional. No instance of a higher number than this
is known to me. Again, it is only among the highest Passeres that
the number nine is found, the Oscines having indifferently nine or
ten. In a good many Oscines, rated as nine-primaried, there are
actually ten, though the outermost is so rudimentary, and even out
of alignment with the developed primaries, that it is not counted
as one of them. Among Oscines, just this difference of one evident
and unquestionable primary more or fewer forms one of the best
distinctions between the families of that suborder. So the tenth
feather in a bird’s wing, counting from the outside, becomes a
crucial test in many cases ; for, if it be last primary, the bird is
one thing ; if it be first secondary, the bird is another. In such
cases the necessity, therefore, of determining exactly which it is
becomes evident. Of course it is always possible to settle the
question by striking at the roots of the remiges and seeing how
many are seated on the pinion ; but this generally involves some
defacing of the specimen, and there is usually an easier way of
determining. Hold the wing half-spread ; then, , in most Oscines,
the primaries come sloping down on one side, and the secondaries
similarly on the other, to form where they meet a reentrant angle
in the general contour of the posterior border of the wing ; the
feather that occupies this notch is the one we are after, and
unluckily it is sometimes last primary, sometimes first secondary.

SEC. Ill

EXTERIOR FARTS OF BIRDS

167

But observe that primaries are, so to speak, self -asserting, emphatic,
italicised, remiges, stiff, strong, and obstinate ; while secondaries are
retiring, whispering, in brevier, limber, weak, and yielding. Their
different character is almost always shown by something in their
shape or texture which the student will soon learn to recognise,
though it cannot well be described. Let him examine Fig. 30,
where h marks the nine primaries of a sparrow’s wing, and s indi-
cates the secondaries j he will see a difference at once. The
primaries express themselves, though with diminishing emphasis, to
the last one ; then the secondaries begin to tell a different tale.
The condition of the first primary, whether spurious or not, is often
of great help in this determination. The first primary is called
“ spurious ” when it is very short — say one-third, or less, as long
as the second, or longest primary. Among Passeres, a spurious first
primary only occurs in certain ten-primaried 0 seines ; whence it is
evident, that to find such short first primary is equivalent to deter-
mining the presence of ten primaries, though not to find it does not
prove there are only nine ; the count should be made in all cases in
which the outer primary is more than one-third as long as the next.
The difference between nine primaries, and ten with the first spuri-
ous, is excellently illustrated among the species of the American
Vireo. Any thrush, nuthatch, titmouse, or creeper shows a spurious
primary to advantage — large enough not to be overlooked, small
enough not to be mistaken.

The Secondaries (Fig. 30, s) are those remiges Avhich are seated
on the forearm (Fig. 27, ^ to C). They vary in number from six
to forty or more. They have the peculiarity of being attached to
one of the bones of the forearm, the ulna. If an ulna be examined
closely, there will be seen a
row of little points showing
the attachment ; such are indi-

cated in Fig. 27 along nl, and showing

in Fifif. 31. The secondaries points of attachment of the secondaries. (Dr. E.

, X W. Shufeidt, U.S.A.)

present no points necessary to

dwell upon here, after what has been said of the primaries. They
are enormously developed in the Argus pheasant, and have curious
shapes in some other exotic birds. They are often long enough to
cover the primaries completely when the wing is closed, as in
grebes ] on the other hand, they are extremely short in the swifts
and humming-birds.

The Tertiaries (Fig. 30, t) are properly the remiges which grow
upon the upper arm, humerus. But such feathers are not very evident
in most birds, and the two or three innermost secondaries, growing
upon the very elbow, and commonly different from the rest in form
or colour, pass under the name of “ tertiaries.” Again, in some

68

GENERAL ORNITHOLOGY

PART II

cases, scapular feathers (Fig. 30, scj?) are called tertiaries, especially
when long or otherwise conspicuous. But there is an evident and
proper distinction. Scapulars belong to the ;pteryla humemlis (see
p. 130) ; while tertiaries, whether seated on the elbow or higher up
the arm, are the innermost remiges of the jpteryla alaris. These
inner remiges are often shortly called tertials ; though the longer
name is more correct, besides being conformable with the names of
the other two series of remiges. Tertiaries often afford good char-
acters for description, in peculiarities of their size, shape, or colour.
Thus it is very common among Fringillidm for these feathers to be
parti-coloured differently from the other remiges. In many birds
they are long and flowing; as in the families Motacillidce. and Alau-
didoi, where they reach about to the end of the primaries when the
wing is closed. Their development is similar in many Scolopacidce.
In such cases, the feather-border of the wing pronounces the letter
W quite strongly — outer lower angle at point of primaries ; middle
upper angle at reentrance between primaries and secondaries ; inner
lower angle at point of tertiaries.

The “ point of the wing ” is at the tip of the longest primary.
It is best expressed when the first primary is longest. Sometimes
the end is so much rounded off, that the midmost primary may be
the longest one, the others being graduated on both sides of this
projecting point. In speaking of the relative lengths of remiges,
we always mean the way in which their tips fall together, not the
actual total lengths of the feathers. Thus a second primary, whose
tip falls opposite the tip of the first one, is said to be of equal length,
though it may actually be longer, being seated higher up on the
pinion. The development of the primaries also furnishes one of the
most important measurements of birds : for the expression “ length
of wing,’' or simply “ the wing,” means the distance from the “ bend
of the wing,” or carpal angle, to the end of the longest primary.
The integument of the wing does not very often develop anything
but feathers. Occasionally

Claws and Spurs are found upon the pinion. Claws have been
already noticed (p. 160). They are properly so called, being horny
growths comparable in every way to those upon the ends of the
toes, like the claws of beasts, or human nails. A spw' (Lat. calcar),
however, is something different, though of the same horny texture,
since it does not terminate a digital phalanx, but is off-set from the
side of the hand. It is exactly like the spur on the leg of a fowl,
which obviously is not a claw. The spur- winged goose {Plectropterus),
pigeon (Didunculus), plovers {Chettusia, etc.), the Ja9anas {Parra),
and the doubly-spurred screamer (Palamedea), afford examples of
such outgrowths. (See Fig. 53 ter.)

SEC. Ill

EXTERIOR PARTS OF BIRDS

169

III. THE TAIL.

Its Bony Basis. — Time was when birds flew about with long,
lizard-like, bony, and fleshy tails, having the feathers inserted in a
row on either side like the hairs of a squirrel’s. But we have
changed all that distichous arrangement since when the Archceopteryx
was steered with such a rudder through the scenes of its Jurassic
life. Now the true separate coccygeal bones are few, generally
about nine in number, and so short and stunted that they do not
project beyond the general plumage — in fact, scarcely beyond the
border of the pelvis. Anteriorly, within the bony basin of the
pelvis, there are several vertebrae, which, fusing together and with
the true sacrum, are termed urosacral or false tail-bones. To these
succeed the true caudal vertebrae, movable upon each other and
upon the urosacrum. The last one of these, abruptly larger than
the rest, and of peculiar shape, bears all the large tail-feathers,
which radiate from it like the blades of a fan. The true caudal
vertebrae collectively form the coccyx (Gr. kokkv^, hokkux, a cuckoo ;
from fancied resemblance of the human tail-bones to a cuckoo’s
bill; Fig. 56, civ); the enlarged terminal one is the wmer (Lat.
vomer, a ploughshare, from its shape ; not to be confused with a
bone of the skull of same name) or pygostyle (Gr. irvyri, puge, rump,
and cttuAos, stulos, a stake, pale ; Fig. ^Q,py). The pygostyle, how-
ever, is a compound bone, consisting of several stunted coccygeal
vertebrae fused in one. The bones are moved by appropriate
muscles, and upon the surface is seated the elaeodochon (p. 129).
The whole bony and muscular affair is familiar to every one as the
“ pope’s nose ” of the Christmas turkey ; it is a bird’s real tail, of
which the feathers are merely appendages. In descriptive ornitho-
logy, however, the anatomical parts are ignored, the word “ tail ”
having reference solely to the feathers. These, like those of the
wings, are of two sorts : the coverts or tectrices, and the rudders or
rectrices (Lat. rectrix, pi. rectrices, a ruler, guider ; because they seem
to steer the bird’s flight) ; corresponding exactly to the coverts and
remiges of the wings. The

Tail-Coverts are the numerous comparatively small and weak
feathers which overlie and underlie the rectrices, covering their
bases and extending a variable distance toward their ends, con-
tributing to the firmness and symmetry of the tail. They pass
smoothly out from the body, by gradual lengthening, there being
seldom, if ever, any obvious outward distinction between them and
feathers of the rump and belly ; but they belong to the pteryla
caudalis (p. 131). The natural division of the coverts is into an
upper and under set {tectrices superior es, tectrices inferiores). The

170

GENERAL ORNITHOLOGY

PART II

inferior coverts are the best distinguished from the general plumage,
the anus generally dividing off these “vent-feathers,” as they are
sometimes called. It is to the bundle of under tail-coverts, behind
the vent, that the term crissum is most properly applied. Neither
set is ever entirely wanting ; but one or the other, particularly the
upper one, may be very short, as in a cormorant, or duck of the
genus Erismatura, exposing the quills almost to their bases. While
the upper coverts are usually shorter and fewer than the under
ones, reaching less than half-way to the end of the tail, they some-
times take on extraordinary development and form the bird’s
chiefest ornament. The gorgeous, iridescent, argus-ejmd train of
the peacock consists of enormous tectrices, not rectrices ; the elegant
plumes of the paradise trogon, Phojwmacrus mocinno, several times
longer than the bird itself, are likewise coverts. Occasionally, a
pair of coverts lengthens and stiffens, and then resembles true tail-
feathers, as in the ptarmigan {Lagopus). The crissal feathers are
more uniform in development; they ordinarily form a compact,
definite bundle, as well shown in a duck, where they reach about to
the end of the tail. In some of the storks, they become plumes
of considerable pretensions known as marabous ; and in the
wonderful humming-bird, Loddigesia miraUlis, the middle pair stiffens
to resemble rectrices and projects far beyond the true tail. The
Reetriees, Rudders, or true tail-feathers, like the remiges or
rowers, are usually stiff, well-pronounced feathers, pennaceous to
the very base of the vexilla, without
^ after-shafts, as a rule, and with the

outer web narrower than the other in

most cases. They are always in

pairs ; that is, there is an equal

number of feathers on the right and

left half of the tail ; and their num-

® ~ her, consequently, is an even one.

The exceptions to this rule are so few
and irregular, and then only among birds with the higher numbers
of rectrices, that such are probably to be regarded as mere anomalies,
from accidental arrest of a feather. They are imbricated over each
other in this wise : the central pair are highest, lying with both
their webs over the next feather on either side, the inner web of
one of these middle feathers indifferently underlying or overlying
that of the other ; all thus successively overlying the next outer
one so that they would form a pyramid were they thick instead of
being so flat. The arrangement is perceived at once in the accom-
panying diagram ; where it will be seen, also, that spreading the
tail is the divergence of a from b, while closing the tail is bringing
a and b together under c. The motion is effected by certain muscles

SEC. Ill

EXTERIOR TARTS OF BIRDS

171

that draw on either side upon the bases of the quills collectively ;
they are the same that pull the whole tail to one side or the other,
acting like the tiller-ropes of a boat’s rudder. The general

Shape of a Reetrix is shown in Fig. 23. Such a feather is
ordinarily straight, somewhat clubbed or oblong, widening a little,
regularly and gradually toward the tip, where it is gently rounded

Fig. 32,— The Lyre-bird of Australia, Menura superba, to show the unique lyrate shape of
the tail.

off. But the departures from such shape, or any that could be
assumed as a standard, are numberless, and in some cases extreme.
In fact, none of a bird’s feathers are more variable than those of
the tail ; it is impossible to specify all the shapes they assume.
While most are straight, some are curved — and the curvature may
be to or from the middle line of the body, in the horizontal plane,
or up and down, in the vertical plane. Some shapes have received

172

GENERAL ORNITHOLOGY

PART II

particular names. A rectrix, broad to the very tip, and there cut
squarely off, is said to be truncate; one such cut obliquely off is
incised, especially when, as often happens, the outline of the cut-off
is concave. A linear rectrix is very narrow, with parallel sides j a
lanceolate one is broader at the base, thence tapering regularly and
gradually to the tip. A notably pointed rectrix is said to be acute ;
when the pointing is produced by abrupt contraction near the tip,
as in most woodpeckers, the feather is acuminate. A very long and
slender, more or less linear feather is called filamentous, as the lateral
pair of a barn-swallow or most sea-swallows. The vanes sometimes
enlarge abruptly at the end, forming a spoon-shaped or spatulate
feather ; or such a spoon may result from narrowing of the vanes
near the end, or their entire absence, as in the “ racket ” of a saw-
bill (Momohts). The vanes are sometimes wavy as if crimped ; Plotus
is a fine example of this. Sometimes the vanes are entirely loosened,
the barbs being remote from' each other, as in the exotic genus Stipi-
turus, and some parts of the wonderful caudal appendage of the male
lyre-bird {Menura superla, Fig. 32). When the rhachis projects
beyond the vanes, the feather is spinose, or better, mucronate (Lat.
mucro, a pricker), as excellently shown in a chimney-swift of the genus
Chcetura. A pair of feathers abruptly extending far beyond the
others are called long-exserted, after the analogous use of the term in
botany. Tail-feathers also differ much in their consistency, from the
softest and weakest, not well distinguished from coverts, to such stiff
and rugged props as the woodpeckers possess. They are downy and
very rudimentary in a few birds, notably all the grebes, Podicipedidce,
which are commonly said to have no tail. The tinamous of South
America {Dromceognathce) are also very closely docked. The

Typical Number of Reetriees is twelve. This holds in the
great majority of birds. It is so uniform throughout the great
group Oscines, that the rare exceptions seem perfectly anomalous.
In the other group of Passeres {Clamatores) it is usually twelve,
sometimes ten. Ten is the rule among Picarice, though many have
twelve, a very few only eight, as in the genus Crotophaga. The
whole of the woodpeckers (Picidce) have apparenthj ten ; but really
twelve, of which the outer one on each side is spurious, very small,
and hidden between the bases of the second and third feathers.
Birds of prey {Baptores) have usually twelve. In pigeons the rule
is twelve or fourteen ; but sixteen are found in some, and twenty
in one case. In birds below these, the number increases directly ;
there are often or usually more than twelve in the grouse family,
and there may be sixteen, eighteen, or twenty, as among American
genera of Tetraonidce^. Wading birds, often having but twelve,
furnish instances of as many as twenty. Those swimming birds
with large well-formed tails, as the Longipennes, and some Anatidce

SEC. Ill

EXTERIOR FARTS OF BIRDS

173

have the fewest, as twelve, sometimes fourteen, rarely sixteen ;
those with short soft tails have the most, as sixteen to twenty-four.
Among the penguins there are thirty-two or more. The Archceo-
pteryx appears to have had forty, — a pair to each free caudal vertebra ;
and this may be considered the prototypic relation between the
bones and feathers of the tail. The

Typical Shape of the Tail, as a whole, is the fan. The modi-
fications of form, however, which are greater and more varied than
those of the wing, are susceptible of better definition, and many of
them have received special names. Taking the simplest case, where
the rectrices are all of the same length, we have what is called the
even^ square, or truncate tail. The other forms depart from this
mainly by shortening or lengthening of certain feathers. A tail
nearly or quite even may have the two central feathers long-exserted,
as seen in the jaegers {Stercorarius), and tropic-birds {Pha'ethon).
The most frequent departure from the even shape results from
gradual shortening of successive rectrices from the middle to the
outer ones. This is called, in general, gradation or graduation (Lat.
gradus, a step) ; such shortening may be to any degree. More
precisely, graduation means shortening of each successive feather to
the same extent, — say, each half an inch shorter than the next ;
but such exactitude is not often expressed. When the feathers
shorten by more and more, we have the true rounded tail, probably
the commonest form among birds ; thus, the gradation between
the middle and next pair may be just appreciable, and then increase
regularly to an inch between the next and the lateral feather. The
opposite gradation, by less and less shortening, gives the wedge-
shaped or cuneate (Lat. cuneus, a wedge) tail ; it is well shown by
the magpie {Pica) in which, as in many other birds, the middle
feathers would be called long-exserted were the rest all as short as
the outer one is. A cuneate tail, especially if the feathers be
narrow and lanceolate, is also called acute, or pointed, as in the
sprig-tailed duck {Dafila) or sharp-tailed grouse (Pedioecetes). The
generic opposite of the gradated is the forked tail; in which the
lateral feathers successively increase in length from the middle to
the outermost pair. The least appreciable forking is called emar-
gination, and a tail thus shaped is said to be emarginate ; when it is
better marked, as, for instance, an inch of forking in a tail six
inches long, the tail is truly forked or furcate (Lat. furca, a fork).
But the degrees of furcation, like those of gradation, are so insen-
sibly varied, that qualified expressions are usual ; as, “ slightly
forked,” “ deeply forked.” Deep furcation is usually accompanied
by more or less narrowing or filamentous elongation of the lateral
pair of rectrices, as in the barn swallows (Hirundo) and most of the
sea-swallows {Sterna). An advisable term to express such an ex-

74

GENERAL ORNITHOLOGY

PART II

treme furcation is forficate (Lat. forfex, scissors), when the depth of
the fork is at least equal to the length of the shortest feathers ; it
occurs among the birds last named, in the species of the tyrannine
genus Milvulus, and elsewhere. Double-forked and double-rounded tails
are not uncommon ; they result from combination of both opposite
gradations, in this way : The middle feathers being of a certain
length, the next two or three pairs progressively increasing in
length, and the rest successively decreasing, the tail is evidently
forked centrally, rounded externally, which is the double-rounded
form, each half of the tail being rounded ; it is shown in the genera
Myiadestes and Anous. Now if with middle feathers as before, the
next pair or two decrease in length, and then the rest increase to
the outermost, we have the double-forked, a common style among
sandpipers, as if each half of the tail were forked. But in such
case, the forking is slight, merely emargination, being little more

Fig. 33. — Diagram of shapes of tail, adc, rounded ; aec, gradate ; aic, cuneate-gradate ; ale,
cuneate ; abc, double-rounded ; feg, square ; fhg, emarginate ; fneog, double-emarginate ; kim,
forked ; kern, deeply forked ; khm, forficate.

than protrusion of the middle pair of feathers in an otherwise lightly
forked tail ; and in the double-rounded form the gradation is seldom
if ever great.

I should also allude to shapes of tail resulting from the relative
positions of the feathers. Prominent among these is the complicate
or folded tail of the barn-yard fowl, and others of the Phasianidee, —
a very familiar but not common form. It is only retained while
the tail is closed and cocked up, — for when it is lowered and spread
in flight it flattens out. The males of some of the African whydah
birds {Viduince) have remarkably large and long tails of somewhat
similar character. The opposite disposition of the feathers is seen
to some extent in crow-blackbirds {Quiscalus), where the lateral
feathers slant upward from the lowermost central pair, like the
sides of a boat from its keel ; this is the scaphoid (Gr. a-Kacj^T], a
boat) or carinate (Lat. carina, a keel) tail. The American “boat-

SEC. Ill

EXTERIOR FARTS OF BIRDS

175

tailed ” grackle has been so named on this account. One of the
most beautiful and wonderful of all the shapes of the tail is illus-
trated by the male of the lyre-bird {Menura suj)erh% Fig. 32), in
which the feathers are anomalous both in shape and in texture, and
the resulting form of the whole is unique. It should be remembered
that, to determine the shape, the tail should be nearly closed ; for
spreading will make a square tail round, an emarginate one square,
etc. I give a diagram of the principal forms (Fig. 33).

IV. THE FEET.

The Hind Limbs, in all birds, are organised for progression — all
can walk, run, or hop on land, though the power to do so is very
slight in some of the lower swimming birds, as loons and grebes,
and certain of the lower perching birds, as hummers, swifts, goat-
suckers, and kingfishers. They are specially fitted for perching on
trees, bushes, and other supports requiring to be grasped, in the
great majorit}^ of birds, as throughout the Passeres, Picarice., Accipitres,
Columbce, and, in fact, many water-birds ; there being few forms,
mainly found among three-toed birds, or those in which the hind
toe is short, weak, and elevated, in which the extremity of the limb
has not decided grasping power. The limb becomes a paddle for
swimming either on or in the water in many cases. In not a few,
as parrots and birds of prey, the foot is serviceable as a hand.
Those kinds of birds which live in trees and bushes habitually
progress, even when on level ground, in a series of hops, or rather
leaps, both feet being moved together : in all the lower birds, how-
ever, the feet move one after the other, as in ordinary walking or
running. The modifications of the hind limb are more numerous,
more diverse, and more important in their bearing on classification,
than those of either bill, wing, or tail ; their study is consequently
a matter of special interest.

Theip Bony Framework (Fig. 34). — Beginning at the hip-joint,
and ending at the extremities of the several toes, the skeleton of
the hind limb consists in the vast majority of adult birds of twenty
bones. This is the typical and nearly the average number ; birds
scarcely ever have more, and the principal lessenings of the number
result from the absence of one or two toes, or a slight reduction in
the number of the joints of some toes, or absence of the knee-cap.
Of the normal twenty, fourteen are bones of the toes ; one is an in-
complete bone connecting the hind toe with the foot ; one is the
knee-cap, and four are the principal bones of the thigh (1), leg (2),
and foot (1). The first or uppermost is the thigh-bone or femur
femur ; adjective, femoral), fm, from hip to knee, ^ to ^ in the
figure. It is a rather short, quite stout, cylindrical bone, enlarged

176

GENERAL ORNITHOLOGY

PART II

above and below. Above it has a globular head, standing off
obliquely from the shaft, received in the acetabulum (Lat. aceta-
bulum, a kind of receptacle) or socket of the hip, and a prominent
shoulder or trochanter, which abuts against the brim of the acetabulum.
Below, it expands into two condyles (Gr. kovSvXos, hondulos, a knob),

Fig. 34. — Bones of a bird’s right hind limb : from a duck, Clangula islandica, f nat. size •
Dr. R.W. Shufeldt, U.S.A. A, hip ; B, knee ; C, heel or ankle-joint, suffrago ; D, bases of toes.’
A to B, thigh or “ second joint” ; B to C, crus, leg proper, “drumstick,” often wrongly called
“ thigh” ; 0 to D, metatarsus, foot proper, corresponding to our instep, or foot from ankle to
bases of toes ; in descriptive ornithology the tarsus; often called “ shank.” From D outward
are the toes or digits, fm, femur ; tb, tibia, principal (inner) bone of leg ; Ji, fibula, lesser
(outer) bone of leg ; mt, principal metatarsal bone, consisting chiefly of three fused metatarsal
bones ; am, accessory metatarsal, bearing li, first or hind toe, with two joints ; 2t, second toe,
with three joints ; 3^, third toe, with four joints ; 4<, fourth toe, with five joints. At 0 there
are in the embryo some small tarsal bones, not shown in the figure, uniting in part with the
tibia, which is therefore a tibio-tarsus, in part with the metatarsus, which is therefore a tarso-
metatarsus; the ankle-joint being therefore between two rows of tarsal bones, not, as it appears
to be, directly between tibia and metatarsus.

for articulation with both the bones it meets at the knee. It is the
same bone as the femur of a quadruped or of man, and corresponds
to the humerus of the wing. In the knee-joint, many or most birds
have a small ossicle, and a few have two such bony nodules, not
shown in this figure, but nearly in the position of the letter B ; it

SEC. Ill

EXTERIOR FARTS OF BIRDS

177

is the knee-pan or knee-cap, patella (Lat. patella). The thigh is the
first segment of the limb ; the next segment is the leg proper, or
crus (Lat. crus, the shin ; adjective, crural), B to C m the figure, or
from knee to heel. This segment is occupied by two bones, the
tibia (Lat. tibia, a tube, trumpet), tb, and fibula (Lat. fibula, a splint,
clasp), fi. Of these the tibia is the principal, larger, inner bone,
running quite to the heel; the fibula is smaller, and (with rare
exceptions, as in some of the penguins) only runs part way down
the outside of the tibia as a slender pointed spike, close pressed
against or even partly fused with the shaft of the tibia. Above, at
the knee, both bones articulate with the femur ; the tibia with both
the femoral condyles, the fibula only with the outer condyle.
Above, the tibia has an irregularly expanded head or cnemial pro-
cess (Or. Kviigr], kneme), which in some birds, as loons, runs high
up in front above the knee-joint. Below, the tibia alone forms the
ankle-joint, C, by articulating with the next bone. For this purpose
it ends in an enlarged trochlear (Gr. rpoyaAia), or pulley-like surface,
presenting a little forward as well as downward, above which, in
many birds, there is a little bony bridge beneath which tendons
passing to the foot are confined. This finishes the leg, consisting
of thigh, A B, and leg proper, B C, bringing us to the ankle-joint
at the heel, C.

Now a bird’s legs, unlike ours, are not separate from the body
from the hip downward ; but, for a variable distance, are enclosed
within the general integument of the body. The freedom of the
limb is greatest among the high perching birds, and especially the
Raptores, which use the feet like hands, and least among the lowest
swimmers. The range of variation, from greatest freedom to most
extensive enclosure of the limb, is from a little above B nearly to C —
the latter in the case of a loon, grebe, or penguin. In no bird is
the knee, B, seen outside the general contour of the plumage : it
must be looked or felt for among the feathers, and in most prepared
skins will not be found at all, the femur having been removed. It
is a point of little practical consequence, though bearing upon the
generalisation just made. The first joint, or bending of the limb,
that appears beyond a bird’s plumage is the heel, or suffrago, C ; and
this is what, in loose popular parlance, is called “knee,” upon the
same erroneous notions that make people call the wrist of a horse’s
fore-leg “ knee.” People also call a bird’s crus or leg proper, B to
C, the “thigh,” and disregard the true thigh altogether. This con-
fusion is inexcusable ; any one, even without the slightest anatomical
knowledge, can tell knee from heel at a glance, whatever their
respective positions relative to the body. Knee is at junction of
thigh and leg proper ; it always bends forward ; heel is at junction
of leg with foot, and always bends backward. This is as true of a

178

GENERAL ORNITHOLOGY

PART II

bird, which is digitigrade, that is, walks on its toes with its heels in
the air, as it is of a man, who is plantigrade that is, walks on the
whole sole of the foot, with the heel down to the ground. In a
carver’s language, the thigh is the “second joint” (from below);
the leg is the “ drumstick ” ; the rest of a fowl’s hind limb does not
usually come to table, having no flesh upon it.

Before proceeding to the next segment of the limb, I must dwell
upon the ankle-joint, situated at the heel, — the point (7, — corre-
sponding to the carpal angle or bend of the wing, (7, in Fig. 27.
There we found, in adult birds, two small carpal bones, or bones of
the wrist proper ; and noted the presence in the embryo of several
other carpals (Fig. 29), which early fuse with the metacarpus. Just
so in the ankle, there are in embryonic life several tarsal bones, or
bones of the tarsus (Lat. tarsus, the ankle) ; all of which, however,
soon disappear, so that there appears to be no tarsus, or collection
of little bones between the tibia and the next segment of the limb,
the metatarsus. An upper tarsal bone, or series of tarsal bones, fuses
with the lower end of the tibia, making this leg-bone really a tihio-
tarsus ; and similarly, a lower bone or set of bones fuses with the
upper end of the metatarsus, making this bone a tarso-metatarsus.
So there are left no free bones in the ankle-joint, which thus appears
to be immediately between the leg-bone and the principal foot-bone ;
but which is nevertheless really between two series of tarsal bones,
the separateness or identity of which has been lost.^

The next segment of the limb, C to D, or the foot proper, is
represented by the principal metatarsal bone, mt. This corresponds
to the human instep or arch of the foot, nearly from the ankle-joint
quite to the roots of the toes. The metatarsal bone, like the meta-
carpal of the hand, which it represents in the foot, is a compound

^ The exact homologues of a bird’s vanishing tarsal bones are still questioned.
Gegenbaur showed the so-called epiphysis or shoe of bone at the foot of the tibia,
and the similar cap of bone on the head of the principal metatarsal bone, to be true
tarsal elements. Morse went further, showing the tibial epiphysis, or upper tarsal
bone of Gegenbaur, to be really two bones, which he held to correspond with the
tibiale and fibulare, or astragalus and calcaneum of mammals ; these subsequently
combining to form the single upper tarsal bone of Gegenbaur, and finally becoming
anchylosed with the tibia to form the bitrochlear condylar surface so character-
istic of the tibia of Aves. The distal tarsal ossicle he believed to be the
centrale of reptiles. Wyman discovered the so-called “ process of the astragalus ” to
have a distinct ossification, and Morse interpreted it as the intermedium of reptiles.
Later views, however, as of Huxley and Parker, limit the tibial epiphysis to the
astragalus alone of mammals. If these opinions be correct, other tarsal elements
(more than one) are to be looked for in the epiphysis of the metatarsus. Whatever
the final determination of these obscure points may be, it is certain that, as said in
the text above, the lower end of a bird’s tibia and the upper end of a bird’s meta-
tarsus include true tarsal elements, just as the upper end of the metacarpus includes
carpal elements ; and that a bird’s ankle-joint is not tibio-tarsal or between leg-bone
and foot-bones, as in mammals, but between proximal and distal series of tarsal bones,
and therefore medio -idixsdl, as in reptiles.

SEC. Ill

EXTERIOR FARTS OF BIRDS

179

one. Besides including the evanescent tarsal element or elements
already specified, it consists of three metatarsal bones consolidated
in one, just as the metacarpal is tripartite. Among recent birds,
the three are partly distinct only in the penguins ; but in all, ex-
cepting ostriches, the original distinction is indicated by three
prongs or stumps at the lower end of the bone, forming as many
articular surfaces for the three anterior toes. The other toe most
birds possess, the hind toe, is hinged upon the metatarsus in a
different way, by means of a small separate metatarsal bone, quite
imperfect ; this is the accessory metatarsal, am. It is situated near
the lower end toward the inner side of the principal metatarsal

Fig. 34 6is. — Diagram of corresponding segments of hind limbs of man, horse, and bird.
The lines 1-11 are isotomes, cutting the limbs into morphologically equal parts, or isomeres.

bone, and is of various shapes and sizes ; it has no true jointing with
the latter, but is simply pressed close upon it, much as the fibula is
applied to the tibia, or partly soldered with it. Above, it is defect-
ive ; below, it bears a good facet for articulation with the hind
toe. In spite of anatomical proprieties, the metatarsal part of a
bird’s foot — from heel to base of toes — from C to D, is in ordinary
descriptive ornithology invariably called “ The Tarsus ” a wrong
name, but one so firmly established that it would be finical and
futile to attempt to substitute the correct name. In the ordinary
attitude of most birds, it is held more or less upright, and seems to
be rather “ leg ” than a part of the “ foot.” It is vulgarly called

i8o

GENERAL ORNITHOLOGY

PART II

“the shank.” These points must be ingrained in the student’s
mind to prevent confusion (Fig. 34 his).

The digits of the foot, or toes., upon which alone most birds walk
or perch, consist of certain numbers of small bones placed end to
end, all jointed upon one another, and the basal or proximal ones
of each toe separately jointed either with the principal or the acces-
sory metatarsal bone. Like those of the fingers, these bones are
called (Lat. phalanx^ a rank or series) or internodes (because

coming between any two joints or nodes of the toes). The further-
most one of each almost invariably bears a nail or claw (unguis).
The phalanges are of various relative lengths, and of a variable
number in the same or different toes. But all these points, being
matters of descriptive ornithology rather than of anatomy proper,
are fully treated beyond, as is also the special horny or leathery
covering of the feet usually existing from the point C outward. We
may here glance at the

Mechanism of these Bones. — The hip is a ball-and-socket joint,
permitting roundabout as well as fore-and-aft movements of the
whole limb, though more restricted than the shoulder-joint. The
knee is usually a strict ginglymus (Gr. yiyyXvfxos, gigglumos, hinge)
or hinge-joint, allowing only backward and forward motion ; and so
constructed that the forward movement of the leg is never carried
beyond a right line with the femur, while the backward is so exten-
sive that the leg may be quite doubled under the thigh. In some
birds there is a slight rotatory motion at the knee, very evident in
certain swimmers, by which the foot is thrown outward, so that the
broad webbed toes may not “ interfere.” The heel or ankle-joint is
a strict hinge ; its bendings are just the reverse of those of the knee ;
for the foot cannot pass back of a right line with the leg, but can
come forward till the toes nearly touch the front of the knee. In
some birds the details of structure are such that, with the assistance
of certain muscles, the foot is locked upon the leg when completely
straightened out, so firmly that some little muscular elfort is required
to overcome the obstacle j birds with this arrangement sleep securely
standing on one leg, which is the design of the mechanism. The
jointing of the toes with the prongs of the metatarsus is peculiar ;
for the articular surfaces are so disposed in a certain obliquity, that
when the toes are brought forwards, at right angles or thereabouts
with the foot, they spread apart from each other automatically in
the action, and the diverging toes of the foot thus opened are
pressed upon the ground or against the water. When the toes are
bent around in the opposite direction, they automatically come to-
gether and lie in a bundle more or less parallel with one another,
besides being each bent or flexed at their several nodes. This
mechanism is best marked in the swimmers, which, for advantageous

SEC. Ill

EXTERIOR FARTS OF BIRDS

i8i

use of their webbed toes, must present a broad surface to the water
in giving the backward stroke, and bring the foot forward with the
toes closed, presenting only an edge to the water, — on the principle
of the feathering of oars in rowing. It is carried to an extreme in
a loon, where, when the foot is closed, the digit marked 2t in the
figure lies below and behind 3t. It is probably least marked in
birds of prey, which give the clutch with their talons spread. The
jointings of the individual phalanges of the toes upon one another
are simple hinges, permitting motion of extension to a right line or
a little beyond in some cases, with very free flexion in the opposite
direction. On the whole, the mechanics of a bird’s foot are less
peculiar than those of the wing, and quite like those of the limbs
of a quadruped.

In ordinary hopping, walking, and running, and in perching as
well, only the toes rest upon or grasp the support, from D to be-
yond, C being more or less vertically over D. Such resting of the
toes is complete for 2t, 3t, M in the figure, or for all the anterior
toes ; but for the hind toe it varies according to the length and
position of that digit, from complete incumbency, like that of the
front toes, to mere touching of the tip of that toe, or not even this :
the hind toe is then sure to be functionless. But many of the
lower birds, such as loons and grebes, cannot stand at all upright on
their toes, but rest with the heel (C) touching the ground ; and in
many such cases the tail furnishes additional support, making a
tripod with the feet, as in the kangaroo. Such birds might be
called plantigrade (Lat. planta, the sole ; gradus, a step) in strict
anatomical conformity with the quadrupeds so designated. The
others are all digitigrade, standing or walking on their toes alone.
But no birds progress on the ends of their toes, or toe-nails, as
hoofed quadrupeds do. A bird’s walking or running is the same as
ours, so far as the ordinary mechanics of the motions are concerned ;
but its so-called “ hopping ” is really leaping, both legs moving at
once. Most birds, down to Columbce, leap when on the ground, a
mode of progression characteristic of the higher orders ; but many
of the more terrestrial Passer es and Accipitres progress by ordinary
walking when on the ground, as is invariably the case with parrots,
pigeons, gallinaceous birds, and all waders and swimmers.

The student need scarcely be reassured that, whatever their modi-
fications, their relative development, motions, and postures, the
several segments of both fore and hind limbs of any vertebrate,
quadruped or biped, feathered or featherless, are fixed in one mor-
phologically identical series, thus : 1, shoulder or hip-joint ; 2, upper
arm or thigh, humerus or femur ; 3, elbow or knee-joint ; 4, fore-
arm or leg proper, radius and ulna or tibia and fibula ; 5, wrist,
bend of wing, carpus, or heel, ankle, tarsus ; 6, hand proper, meta-

GENERAL ORNITHOLOGY

PART II

182

carpus, or foot proper, metatarsus ; 7, digits with their phalanges,
of hand or foot, fingers or toes. Observe the improper popular
naming of these parts, in the case of the hind limb, whereby 1, 2, 3,
are not generally counted ; 4 is miscalled “ thigh ” ; 5 is miscalled
“knee”; 6 is miscalled “leg” or “shank”; 7 is miscalled “foot.”
Observe also that in descriptive ornithology 6 is “ the tarsus^

The Plumage of the Leg and Foot varies within wide limits.
In general, the leg is feathered to the heel, C, and the rest of the
limb is bare of feathers. The thigh is always feathered, as part of
the body plumage {pteryla femoralis). The crus or leg proper (thigh
of vulgar language, B to C) is feathered in nearly all the higher
birds, and in swimming birds without exception ; in the loons, the
feathering even extends on the heel-joint. It is among the walking
and especially the wading birds that the crus is most extensively
denuded ; it may be naked half-way up to the knee. A few waders
— among British birds, chiefly in the snipe family — have the crus

Fig, 35. — Feathered tarsus of the prairie-hen, Cupidonia cupido. Nat. size ; from life by Coues.

apparently clothed to the heel-joint ; but this is due, in most if not
all cases, to the length of the feathers, for probably in none of them
does the pteryla cruralis itself extend to the joint. Crural feathers
are nearly always short and inconspicuous ; but sometimes long
and flowing, as in the “ flags ” of most hawks, and in the American
tree-cuckoos (Coccyzus). The tarsus (I now and hereafter use the
term in its ordinary acceptation — C to D in Fig. 34 ; trs in Fig. 36)
in the vast majority of birds is entirely naked, being provided with
a horny or leathery sheath of integument like that covering the
bill. Such is its condition in the Passeres and Pkarice (with few
exceptions, as among swifts and goatsuckers) ; in the waders with-
out exception, and in nearly all swimmers (the frigate-bird, Tachy-
petes, has a slight feathering). The Raptores and Gallince furnish the
most feathered tarsi. Thus, feathered tarsi is the rule among owls
(Striges) ; frequent, either partial or complete, in hawks and eagles,
as in Aquila, Archibuteo, Falco, Buteo, etc. All British grouse, and
perhaps all true grouse, have the tarsus more or less feathered (Fig.

SEC. Ill

EXTERIOR FARTS OF BIRDS

183

35). The toes themselves are feathered in a few birds, as several of
the owls, and all the ptarmigans {Lagopus). Partial feathering of
the tarsus is often continued downward, to the toes or upon them,
by sparse modified feathers in the form of bristles j as is well shown
in the barn-owl (Fig. 47). When incomplete, the feathering is
generally wanting behind and below, and it is almost invariably
continuous above with the crural plumage. But in that spirit of
perversity in which birds delight to prove every rule we establish
by furnishing exceptions, the tarsus is sometimes partly feathered
discontinuously. A curious example of this is afforded by the bank-
swallow, Cotile riparia, with its little tuft of feathers at the base of
the hind toe ; and some varieties of the barn-yard fowl sprout mon-
strous leggings of feathers from the side of the tarsus.

• The Length of Leg, relatively to the size of the bird, is ex-
tremely variable ; a thrush or sparrow probably represents about
average proportions of the limb. The shortest-legged bird known
is probably the frigate-pelican (Tachypetes) ; which, though a yard
long, more or less, has a tibia not half as long as the skull, and a
tarsus under an inch. The leg is very short in many Picarian birds,
as hummers, swifts, goatsuckers, kingfishers, trogons, etc., in most
of which it scarcely serves at all for progression. Among Passeres,
the swallows resemble swifts in shortness of their hind limbs. It
is pretty short likewise in many zygodactyl, yoke-toed, or scansorial
birds, as woodpeckers, cuckoos, and parrots. In most swimming
birds the limb may also be called short, especially in its femoral
and tarsal segments ; while the broad-webbed toes are comparatively
longer. The leg lengthens in the lower perching birds, as many
hawks and some of the terrestrial pigeons ; it is still longer among
walkers proper, such as the gallinaceous birds, and reaches its
maximum among the waders, especially the larger ones, such as
cranes, herons, ibises, storks, and flamingoes; among all of which
it is correlated with extension of the neck. Probably the longest-
legged of all birds for its size is the stilt (Himantopus). Taking
the tarsus alone as an index of length of the whole limb, this is in
the frigate under one-thirty-sixth of the bird’s length ; a flamingo,
four feet long, has a tarsus a foot long ; a stilt, fourteen inches
long, one of four inches; so that the maximum and minimum
lengths of tarsus are nearly thirty and under three per cent of a
bird’s whole length.

The Horny Integument of the Foot requires particular atten-
tion. That part of the limb which is devoid of feathers is covered,
like the bill, by a hardened, thickened, modified integument, vary-
ing in texture from horny to leathery. This sheath is called the
podotheca (Gr. ttov?, ttoSo?, pous, podos, foot, and O'ijkt], theke, sheath).
It is more corneous in land birds, and in water birds more leathery ;

184

GENERAL ORNITHOLOGY

PART II

this general distinction has hut few exceptions. The perfectly
horny envelope is tight, and immovably fixed or nearly so, while
the skinny styles of sheath are looser, and may usually be slipped
about a little. The integument may differ on different parts of
the same leg, and in fact generally does so to some extent. Unlike
the sheath of the bill, the podotheca is never simple and continuous,
being divided and subdivided in various ways. The lower part of
the crus, when naked, and the tarsus and toes, always have their
integument cut up into scales, plates, tubercles, and other special
formations, which have received particular names. The manner
and character of such divisions are often of the utmost consequence
in classification, especially among the higher birds, since they are
quite significant of genera, families, and even some larger groups.

StclL

Fig, 37. Scutel- 38. — a, Reticulate tarsus of a

late lamini plantar plover. Nat. size, h, Scutellate and
Fig. 36. — Booted laminiplantar tarsus of a cat- reticulate tarsus of a pigeon. Nat.
tarsus of a robin (Tardus migrratoriws). bird (Mimus caro- size.

Nat. size. linensis). Nat. size.

The commonest division of the podotheca is into scales or scutella
(Lat. scutellum^ a little shield; pi. scutella^ not scutellce as often
written); Figs. 37 and 38, h. These are generally of large com-
parative size, arranged in definite vertical series up and down the
tarsus and along the toes, and apt to be somewhat imbricated, or
fixed shingle -wise, the lower edge of one overlapping the upper
edge of the next. The great majority of birds have such scutella.
They oftenest occur on the front of the tarsus (or acrotarsium, corre-
sponding to our “ instep ”), and almost invariably on the tops of the
toes (collectively called acropodium) ; frequently also on the sides
and back (planta) of the tarsus ; not so often on the crus ; and
rarely if ever on the sides and under surfaces of the toes. A tarsus
so disposed as to its podotheca is said to be scutellate, — scutellate

SEC. Ill

EXTERIOR FARTS OF BIRDS

185

before (Fig. 37), or behind, or both, as the case may be. The term
is equally applicable to the acropodium, but is not so often used
because scutellation of the upper sides of the toes is so universal as
to be taken for granted unless the contrary condition is expressly
said. The most notorious case of the Osdm podotheca (Figs. 36, 37),
characterising that great group of birds, is given in the next
paragraph.

Plates, or reticulations (Lat. reticulum, a web ; Fig. 38, a) result
from the cutting up of the envelope in various ways by cross lines.
Plates are of various shapes and sizes, and grade usually into true
scutella, from which, however, they are generally distinguished by
being smaller, or of irregular contour, or not in definite rows, or
lacking the appearance of imbrication; but there is no positive
distinction. They are oftenest hexagonal (six-sided), a form best
adapted to close packing, as shown very perfectly in the cells of
the honey-bee’s comb ; but they may have fewer sides, or be poly-
gonal (many-sided) or even circular ; when crowded in one direction
and loosened in another the shape tends to be oval or even linear.
A leg so furnished is said to be reticulate : the reticulation may be
entire, or be associated with scutellation, as often happens (Fig. 38, h).
A particular case of reticulation is called granulation (Lat. granum,
a grain), when the plates become elevated into little tubercles,
roughened or not. Such a leg is said to be granular, granulated or
rugose: it is well shown by parrots, and the osprey (Pandion).
When the harder sorts of scales or plates are roughened without
obvious elevation, the leg is said to be scabrous or scarious (Lat.
scabrum, a scab). But scabrous is also said of the under surfaces of
the toes, when these develop special pads, or wart-like bulbs (called
tylari) ; as is well shown in many hawks. The softer sorts of legs,
and especially the webs of swimming birds, are often marked cross-
wise or cancellated with a latticework of lines, these, however, not
being strong enough to produce plates ; it is more like the lines
seen on our palms and finger-tips. The plates of a part of the leg
occasionally develop into actual serrations ; as witnessed along the
hinder edge of a grebe’s tarsus. When an unfeathered tarsus shows
no divisions of the podotheca in front (along the acrotarsium), or
only two or three scales close by the toes, it is said to be booted or
gr eared ; and such a podotheca is holothecal (Gr. 6A0?, holos, whole,
entire, and O'qKrj ; Fig. 36). The generic opposite is schizothecal
(Gr. crxtCo), I cleave), whether by scutellation, or reticulation, or in
any other way the integument may be cut up. A booted or holo-
thecal tarsus chiefly occurs in the higher Oscines, and is supposed
by ornithologists to indicate the highest type of bird structure. It
is, however, found in a few water birds, as Wilson’s stormy petrel
and other species of Oceanites. It is not a common modification.

i86

GENERAL ORNITHOLOGY

PART II

Exceptions aside, it only occurs in connection with an equally
particular condition of the sides and hack of the tarsus, or ]planta.
In almost all Oscine Passeres {Alaudidce are an exception), which
constitute the great bulk of the large order Passeres^ the planta is
covered with one pair of plates or lamince, one on each side, meeting
behind in a sharp ridge; a condition called laminijplantar, in dis-
tinction from the opposite, scutellijplantar, state of the parts. A
holothecal podotheca only occurs in connection with the lamini-
plantar condition, the combination resulting in the perfect “ boot.”
Among British birds it is exhibited by the following genera :
Turdus, Cinclus, Saxicola, Pegulus, Cyanecula, Phyllosco][)us ; and even
birds of these genera, when young, show scutella which disappear
with age by progressive fusion of the acrotarsial podotheca. (Com-
pare Figs. 36, 37.)

The CPUS, when bare of feathers below, may, like the tarsus, be
scutellate or reticulate before or behind, or both ; such divisions of
the crural integument being commonly seen in long-legged wading
birds. Or, again, this integument may be loose, softish, and
movable, not obviously divided, and passing directly into ordinary
skin.

The Tarsus, in general, may be called subcylindrical : it is often
quite circular in cross-section ; generally thicker from before back-
ward, and only rarely wider from one side to the other than in the
opposite direction ; but such a shape as this last is exhibited by the
penguins. When the transverse thinness is noticeable, the tarsus is
said to be compressed ; and such compression is very great in a loon,
in which the tarsus is almost like a knife-blade. Quite cylindrical
tarsi occur chiefly when there are similar scales or plates before and
behind, as happens in the larks {Alaudidce) ; they are rare among
land birds, common among waders. Those swimming birds which
have a very thin skinny podotheca are apt to show traces of the
four-sidedness of the metatarsal bone. The tarsus in the vast
majority of land birds is seen on close inspection to be somewhat
oval or drop-shaped on cross-section, — gently rounded in front,
more compressed laterally, and sharp-ridged behind. This results
from the laminiplantation described above, and is equally well ex-
hibited by most passerine birds, whether they have booted or
anteriorly scutellate tarsi. The line of union of anterior scutella
with posterolateral plates on the sides of the tarsus is generally in a
straight vertical line, — either a mere line of flush union, or a ridge,
or oftener a groove (well seen in the crows), which may or may not
be filled in with a few small narrow plates. In Clamatorial Passeres
the tarsus is enveloped in a scroll-like podotheca of irregularly
arranged plates, the edges of the scroll meeting along the inner side
of the tarsus. — But the full consideration of special states of the

SEC. Ill

EXTERIOR TARTS OF BIRDS

187

tarsal envelope, however important and interesting, would be part
of a systematic treatise on ornithology, rather than of an outline
sketch like this.

The Normal Number of Toes (individually, digiti ; collectively,
podium) is four : there are never more. There are two in the ostrich
alone, in which both inner and hind toe are wanting. There are
three in all the other struthious birds
{Rheidce, Casuariidce), exceigting Apteryx,

which has four. There are likewise pW

three, the hind toe being suppressed, in

the tinamine genera Calodromas and

Tinamotis {Dromceognathce) ; throughout

the auk family {Adcidce) ; in the petrel

genus Pelecandides ; apparently in the

albatrosses {Diomedeince ; in these,

however, there is a rudiment of the

hind toe) ; usually in the gull genus

• in thp flamino-n o-Annc; Pluv'nirn- 39.— Tridactyle foot of sand-

JXlSSw f in tne namin^O genus TUCeniCO erling, Calidris arenarta ; nat. size.

parr a ; throughout the bustard family

{Otididce), and among various related forms, as CEdicnemus,
Hcematopus, Himantopus, Esacus^ Cursorius ; in the plovers (Chara-
driidoe), excepting Squatarola ; in the sandpiper genus Calidris ; and
in the bush-quails (Tiirnicidce), excepting Fedionomus. In higher
birds three toes are a rare anomaly, only known to occur in three
genera of woodpeckers {Pico'ides, Sasia, and Tiga), and in one galbu-
line genus {Jacamaralcyon)^ by loss of the hind toe ; in two genera
of kingfishers ifieyx and Alcyone), by suppression of the inner front
toe j and in the passerine genus Cholornis, by defect of the outer
front toe. Birds with two toes are said to be didactyl ; with three,
tridactyl ; with four, tetradachjl. In the vast majority of cases birds
have three toes in front and one behind. Occasionally either the
hind toe or the outermost front toe is versatile, that is, susceptible of
being turned either way. Such is the condition of the outer front
toe in most owls (Striges), and in the osprey {Pandion). There is no
case of true versatility of the hind toe among North American birds ;
but several cases of its stationary somewhat lateral position, as in
goatsuckers (Caprimulgidce), some of the swifts (Cypselidce), the loons
{Colymhidce), and all the totipalmate swimmers (Steganopodes). The
rarest of all conditions (seen in some Cypselidce, and the African
Coliidce) is that in which all four toes are turned forward. The
arrangement of toes in pairs, two before and two behind, is quite
common, being characteristic of scansorial birds and some others, as
all the parrots and woodpeckers, cuckoos, trogons, etc. Such
arrangement is called zygodactyl or zygodactylous (Gr. C^yov, zugon,
a yoke ; SaKrvXos, daUulos, a digit) ; and birds exhibiting it are said

GENERAL ORNITHOLOGY

PART II

to be yoke-toed (Fig. 45). In all yoke-toed birds, excepting the
trogons, it is the outer anterior toe which is reversed ; in trogons,
the inner one ; the latter are called heterodactylous. In nearly every
three-toed bird, all three toes are anterior ; an exception is in the
genus Picoides, where the true hind toe is wanting, the outer anterior
one being reversed as usual in zygodactyls. No bird has more toes
behind than in front. Birds’ toes, and their respective joints, are
Numbered, in a certain definite order, as follows (see Figs. 34, 36):
hind ioQ= first toe, 1^/ inner anterior tOQ = second toQ, 2t ; middle
anterior toe = third toe, 3t ; outer anterior toe = fourth toe, 4^.
Such identification of It, 2t, 3t, it applies to the ordinary case of
three toes in front and one behind. But, obviously, it holds good
for any other arrangement of the toes, if we only know which one
is changed in position, — a thing always easy to learn, as we shall
see at once. In birds with the hind toe reversed, bringing all four in
front, the same order is evident, though then It is the inner
anterior, 2t the next, etc. ; for it always happens, when a hind toe
turns forward, that it turns on the inner side of the foot. Similarly,
in yoke-toed birds (excepting Trogonidoe), it is the outer anterior
which is turned backward, as above said ; then, evidently, inner
hind toe = l^/ inner front toe = 2t; outer front toe = 3^; outer
hind toe = it. In Trogonidoe, with inner front toe reversed, the
correction of the formula is easily made. Moreover, when the
number of toes decreases from four to three or two, the digits are
almost always reduced in the same order : thus, in three-toed birds,
1^ is the missing one; in the two-toed ostrich. It and 2t are gone.
Exceptions to this generalisation are afforded by two exotic genera
of kingfishers, Ceyx and Alcyone, in which 2t is defective ; and by
the anomalous passerine Cholornis of China, in which it is in like
case. The rule is proved by the

Number of Phalanges, or joints, of the digits. The constancy
of the joints in birds’ toes is remarkable, — it is one of the strongest
expressions of the highly monomorphic character of Aves. In cdl
birds, excepting Procellariidce, \t when present has two joints (not
counting, of course, the accessory metatarsal). In all birds, 2t
when present has three joints. In nearly all birds, 3^ has four
joints. In nearly all birds, it has five joints. Thus, any digit has
one more joint than the number of itself. (See Fig. 34, where the
digits and the phalanges are numbered.) The exceptions to this
regularity consist in the lessening of the number of joints of 1^ or
2>t by one, and of it by one or two. So when the joints do not
run 2, 3, 4, 5, for toes 1 to 4, they run either I, 3, 4, 5, or
2, 3, 4, 4, or 2, 3, 3, 3. (These statements do not regard the
anomalous cases of Ceyx, Alcyone, and Cholornis — see above.) This
variability is nearly confined to certain Picarian birds : examples

SEC. Ill

EXTERIOR FARTS OF BIRDS

1

Fig. 40.— Pha-
langes of Cypse-

of it are in certain genera of Cypselince, Fig. 40, where the ratio is
2, 3, 3, 3, of CaprimulgincB, Fig. 41, where it is 2, 3, 4, 4 ; and the
petrel family, with 1, 3, 4, 5. Such admirable conservatism enables
us to tell what toes are missing in any case, or what ones are out of
the regular position. Thus, in Picoides, the hind toe,
apparently 1/, is known to be 4/, because it is five-
jointed ; in a trogon, the inner hind toe is 2^, being
three-jointed; in the ostrich, with only two toes,

M and At are seen to be preserved, because they
are respectively four- and five- jointed. Besides this

interesting numerical ratio, the phalanges have other
inter-relations of some consequence in classification,
resulting from their comparative lengths. In some
families of birds, one or more of the lasal or proximal 3, 3, s.

’ 1C 1 7 7 oclater.

phalanges (those next to the foot — opposed to distal,
or those at the ends of the digits) of the front toes are extremely
short, being mere nodules of bone (Fig. 40) ; in other and more
frequent cases, they are the longest of all, as in Figs. 34, 41.
On the whole, they generally decrease in length from proximal to
distal extremity, and the last one of any toe is quite small, serving
merely as a core to the claw. The difference in the lengths of the
several phalanges, like that of the digits themselves, makes the
toes more efficient in grasping, since they thereby
clasp more perfectly upon an irregular object. The
design and the principle are the same as seen in
the human hand, in which model instrument the
digits and their respective joints are all of different
lengths.

The Position of the Digits, other than in re-
spect to their direction, is important. In all the birds
front toes are inserted on the metatarsus on the same
level, or so nearly in one horizontal plane that the
difference is not notable. The same may be said of
Fig. 41. — Pha- the hind toes when they are a pair, as in zygodactyle
g^nffoot, birds. But the hind toe, or hallux, as it is called, when

After sciater. present and single, varies remarkably in position

with reference to the front toes ; and this matter requires special
notice, as it is important in classification. The insertion of this
digit varies from the very bottom of the tarsus (nietatarsus), where it
is on a level with the front toes, to some distance up that bone.
When the hallux is flush with the bases of the other toes, so that its
whole length is on the ground, it is said to be incumbent. When
just so much raised that its tip only touches the ground, it is called
insistent. When inserted so high up that it does not reach the
ground, it is termed remote (cmotus) or elevated. But as the precise

190

GENERAL ORNITHOLOGY

PART II

position varies insensibly, so that the foregoing distinctions are not
readily perceived, it is practically best to recognise only two of these
three conditions, saying simply “hind toe elevated,” when it is
inserted fairly above the rest, and “hind toe not elevated,” when
its insertion is flush with that of the other toes. In round terms :
it is characteristic of all insessorial (Lat. insedo, I sit upon) or perch-
ing birds to have the hind toe down ; of all other birds to have it
UP (when present). The exceptions to the first of these statements
are extremely rare.

The Hallux has other Notable Characters. — It is free and
simple, in the vast majority of birds : in all insessorial birds,
nearly all cursorial (Lat. cursor, a courser), and most natatorial
(Lat. natator, a swimmer) forms. Its length, claw included, may
equal or surpass that of the longest anterior toe ; and generally
exceeds that of one or two of these. It is always longest when
incumbent; when thus down on a level with the rest it also
acquires its greatest mobility and functional efficiency. In most
Passeres it has a special muscle for independent movement, so that
it may be perfectly apposable to the other toes collectively, just as
our thumb may be brought against the tip of any finger. In
general, it shortens as it rises on the metatarsus ; and probably in
no bird in which it is truly elevated is it as long as the shortest
anterior toe. It is short, barely touching the ground, in most
wading birds ; shorter still in some swimmers, as the gulls, where it
is probably functionless ; it is incomplete in one genus of gulls
(Rissa), where it bears no perfect claw ; it has only one phalanx and
is represented only by a short immovable claw in the petrels (Pro-
cellariidcB) ; it disappears in the birds named in the last paragraph
but two al30ve, and in some others. It is never actually soldered
with any other toe for any noticeable distance ; but it is webbed to
the base of the inner toe in the loons (Colymhus), and to the whole
length of that toe in all the Steganopodes (Fig. 52). It may also be
independently webbed ; that is, be provided with a separate flap or
lobe of free membrane. This lobation of the hallux is seen in all
our sea-ducks and mergansers {Fuligulince and Merginoe), and in all
the truly lobe-footed birds, as coots (Fulica), grebes (Podicipedidce),
and phalaropes (Phalaropodidce). The modes of union of the
anterior toes with one another may be finally considered under the
head of the

Three leading Modifications of the Avian Foot. — Birds’ feet
are modelled, on the whole, upon one or another of three plans,
furnishing as many types of structure ; which types, though they run
into one another, and each is variously modified, may readily be
appreciated. These plans are the perching or insessorial; the
walking or wading, cursorial or grallatorial ; and the swimming or

SEC. Ill

EXTERIOR FARTS OF BIRDS

191

natatorial — in fact, so well distinguished are they, that carinate birds
have even been primarily divided into groups corresponding to these
three evidences of physiological adaptation of the structure of the
Avian pes. Independently of the number and position of the digits,
the plans are pretty well indicated by the method of union of the
toes, or their entire lack of union. 1. The insessorial ty^e. (a) In
order to make a foot the most of a hand, that is, to fit it best for
that grasping function which the perching of birds upon trees and
bushes requires, it is requisite that the digits should be as free and
movable as possible, and that the hind one should be perfectly
apposable to the others. Compare the human hand, for example,
with the foot, and observe the perfection secured by the entire
freedom of the fingers, and especially the appositeness of the thumb.
In the most accomplished insessorial foot, the front toes are cleft to
the base, or only coherent to a very slight extent ; the hind toe is
completely incumbent, and as
long and flexible as the rest. The
thrushes probably show

as complete cleavage as is ever
seen, practically as much as that
of the human fingers ; the cleft
between the inner and middle toe
being to the very base, while the
outer is only joined to the
middle for about the length of
its own basal joint. This is
the typical passerine foot (Figs.

36, 37, 42, 43). There may ^ ^ i ^

’ ’ ’ ' . Figs. 42, 43.— Typical passerine feet. (The

be somewhat more cohesion right-hand fig. is Plectrophanes lapponicus, nat.

of the toes at base, as in the

wrens, titmice, creepers, etc., without, however, obscuring the true
passerine character. Besides the typical passerine, there are several
other modifications of the insessorial foot, (b) Thus a kingfisher
shows what is called a syndactyl or syngnesious (Gr. crvv, sun, together ;
yv^o-to?, gnesios, relating to way of birth ; Fig. 44) foot where the outer
and middle toes cohere for most of their extent and have a broad sole
in common. It is a degradation of the insessorial foot, and not a
common one either ; seen in those perching birds which scarcely use
their feet for progression, but simply for sitting motionless, (c) The
zygodactyl or yoke -toed modification has been sufficiently noted
(Fig. 45). It was formerly made much of, as a scansorial or climbing
type of foot, and an absurd “ order ” of birds has been called Scan-
sores. But many of the zygodactyl birds do not climb, as the
cuckoos ; while the most nimble and adroit of climbers, such as
nuthatches and creepers, retain a typically passerine foot. The

192

GENERAL ORNITHOLOGY

PART II

scansorial is simply one modification of the insessorial plan, and
has little classificatory significance — no more than that attaching to
the particular condition of the insessorial foot (t?) which results from
elevation or versatility of the hind toe, as in some Cypselidce
Ca^primulgidce. This is an abnormality which has received no

Fig. 44. — Syn-

dactyle foot of king- Fig. 45.— Zygodactyle foot of a woodpecker, Hylotomus

fisher, nat. size. pileatus, nat. size. From nature by Coues.

special name ; it is generally associated with some little webbing
of the anterior toes at base, which is a departure from the true
insessorial plan, or with abnormal reduction of the phalanges of
the third and fourth toes, as explained above (Figs. 40, 41). (e)

The raptorial is another modification of the insessorial foot. It is

Fig. 46. — Kaptorial foot of a hawk, Accipiter cooperi, nat. size. From nature by Coues.

advantageous to a bird of prey to be able to spread the toes as
widely as possible, that the talons may seize the prey like a set of
grappling irons ; and accordingly the toes are widely divergent from
each other, the outer one in the owls and a few hawks being quite
versatile. In a foot of raptorial character, the toes are cleft pro-

SEC. Ill

EXTERIOR FARTS OF BIRDS

193

foundly, or, if united at base, it is by movable webbing; the claws
are immensely developed, and the under surfaces of the toes are
scabrous or bulbous for greater security of the object grasped. Any
hawk or owl or old-world vulture exhibits the raptorial or inses-
sorial foot (Figs. 46, 47). 2. The cursorial or grallatorial type. The

gist of this plan lies in the decrease or entire loss of the grasping
function, and in the elevation, reduction in length, or loss of the
hind toe ; the foot is a good foot, but nothing of a hand. The
columbine birds, which are partly terrestrial, partly arboreal, ex-
hibit the transition from the perching to the gradient foot, in some
reduction of the hind toe, which is nevertheless in most cases still
on the same level as the rest (Fig. 38, h). In the gallinaceous or
rasorial (Lat. rasor, a scraper) birds, which are essentially terrestrial,
and noted for their habit of scratching the ground for food, the

Fig, 47.— Raptorial foot of an owl, Aluco flammeus, nat, size. From nature by Coues.

hind toe is decidedly elevated and shortened in almost all of the
families (Fig. 35). Such reduction and uplifting of the hallux is
carried to an extreme in most of the waders, or Grallatores, in many
of which this toe disappears (Figs. 38 a, 39). It is scarcely practic-
able to recognise special modifications of such gradient or gralla-
torial feet, since they merge insensibly into one another. The herons,
which are the most arboricole of the waders, exhibit a reversion to
the insessorial type, in the length and incumbency of the hallux.
The mode of union of the front toes of the walkers and waders is
somewhat characteristic. The toes are either cleft quite to the
base, or there joined by small webs ; probably never actually
coherent. Such basal webbing of the toes is called semipalmation
(“ half- webbing ”). It is actually the same thing that occurs in
many birds of prey, in most gallinaceous birds, etc. ; the term is
mostly restricted, in descriptive ornithology, to those wading birds,
or Grallatores, in which it occurs. Such basal webs generally run out
O

194

GENERAL ORNITHOLOGY

PART II

to the end of the first, or along part of the second, phalanx of the
toes ; usually farther between the outer and middle than between
the middle and inner toes. Such a foot is well illustrated by the
semipalmated plover JEgialites semipalmatus), semipalmated sand-
piper {Ereunetes pusillus. Fig. 48), and willet {Symphemia semipalmata,
Fig. 49). In a few wading birds, as the avocet and flamingo, the
webs extend to the ends of the toes. This introduces us at once to
the third main modification of the foot. 3. The natatorial type. Here
the foot is transformed into a swimming implement, usually with
much if not entire abrogation of its function as a hand. Swimming
birds, with few exceptions, are bad walkers, and few of them are
perchers. The swimming type is presented under two principal
modifications : — {a.) In the palmate or ordinary webbed foot, all
the front toes are united by ample webs (Fig. 50). The palmation

is usually complete, extending to the ends of the toes j but one or
both webs may be so deeply incised, that is, cut away, that the pal-
mation is practically reduced to semipalmation, as in terns of the
genus Hydrochelidon (Fig. 51). The totipalmate is a special case of
palmation, in which all four toes are webbed ; this characterises the
whole order Steganopodes (Fig. 52). {h.) In the lohate foot, a paddle

results not from connecting webs, but from a series of lohes or flaps
along the sides of the individual toes ; as in the coots, grebes,
phalaropes, and sun- birds {Heliornithidce.) Lobation is usually
associated with semipalmation, as is well seen in the grebes
{Podicipedidce). In the phalaropes {Phalaropodidce, Fig. 53 Us) loba-
tion is present as a modification of a foot otherwise quite cursorial.
The most emphatic cases of lobation are those in which each joint of
the toes has its own flap, with a free convex border; the mem-
branes as a whole therefore present a scolloped outline (Figs. 53, 53
Us). Such lobes are merely a development of certain marginal

Fig. 49. —Semipal-
mated bases of toes
of SympJiemia ; nat.
size.

Fig. 48.— Semipalma-
tion in Ereunetes ; nat.
size.

Fig. 50. — Palmate foot of a tern,
Sterna forsteri ; nat. size.

SEC. Ill

EXTERIOR FARTS OF BIRDS

195

fringes or processes exhibited by many non-lobate or non-palmate
birds. Thus, if the foot of some of the gallinules be examined in a
fresh state, the toes will be seen to have a narrow membranous
margin running the whole length. The same thing is evident in

Fig. 51.— Incised pal-
mation of Hydrochelidon
lariformis ; nat. size.

Fig. 52. — Totipalmate foot of a peli-
can; reduced.

a great many waders, and on the free borders of the inner and
outer toes of web-footed birds. In the grouse family (Tetraonidce)
marginal fringes are very conspicuous ; there being a great develop-
ment of hard horny substance, fringed into a series of sharp teeth
or pectinations (Fig. 35). These formations appear to be deciduous.

Fig. 53- his. — Lobate foot of phalarope,
Fig. 53. — Lobate foot of a coot; reduced. Lohipes liyperhoreus ; nat. size.

that is, to fall off periodically, like parts of the claws of some
quadrupeds (lemmings).

Claws and Spurs. — With rare anomalous exceptions, as in the
case of an imperfect hind toe, every digit terminates in a complete
clan). The general shape is remarkably constant in the class ;

96

GENERAL ORNITHOLOGY

PART II

variations being rather in degree than in kind. A cat’s claw is about
the usual shape : it is compressed, arched, acute. The great talons of
a bird of prey are only an enlargement of the typical shape ; and,
in fact, they are scarcely longer, more curved, or more acute, than
those of a delicate canary bird ; they are simply stouter. The
claws of scansorial birds are very acute and much curved, as well
as quite large. The under surface of the claw is generally exca-
vated, so that the transverse section, as well as the lengthwise out-
line below, is concave, and the under surface is bounded on either
side by a sharp edge. One of these edges, particularly the inner
edge of the middle claw, is expanded or dilated in a great many
birds ; in some it becomes a perfect comb, having a regular series of
teeth. This pectination (Lat. pecten, a comb), as it is called, only
occurs on the inner edge of the middle claw. It is beautifully
shown by all the true herons (Ardeidce) ; by the goatsuckers

Fig. 53 ter. — Foot of Parra gymnostoma, nat. size, showing the long, straight claws. (From
Ridgway Mus. The spurred wing of the same bird is also shown. See p. 168.)

{Gaprimulgidce, Fig. 41); by the frigate pelican {Tachypetes) and
imperfectly by the barn-owl (Aluco flammeus). It is supposed to be
used for freeing parts of the plumage that cannot be reached by
the bill from parasites ; but this is very questionable, seeing that some
of the shortest-legged birds, which cannot possibly reach much of
the plumage with the comb, possess that instrument. Claws are
more obtuse among the lower birds than in the insessorial and scan-
sorial groups, as the columbine and gallinaceous (rasorial) orders,
and most natatorial families. Obtuseness is generally associated
with flatness or depression ; for in proportion as a claw becomes
less acute, so does it lose its arcuation, as a rule. This is well
illustrated by Wilson’s petrel (Oceanites oceanicus), as compared with
others of the same family. Such condition is carried to an extreme
in the grebes {Podicipedidce), the claws of which birds resemble
human finger-nails. Otherwise, deviations from curvature, without
loss of acuteness, are chiefly exhibited by the hind claw of many

SEC. IV

ANATOMY OF BIRDS

197

terrestrial Passeres, as in the whole family Alaudidce (larks), and
some of the finches (Fringillidce), as the species of “ long-spur ”
{Centrophanes). But all the claws are straight, sharp, and pro-
digiously long, in birds of the genus Parra (Fig. 53 ter)-, these
ja^an^s being enabled to run lightly over the floating leaves of
aquatic plants by such increase in the spread of their toes. Claws
are also variously carinate or ridged, sulcate or grooved. In a few
cases they are rounded underneath, so as to be nearly circular in
cross-section, as is the Case with those of the osprey (Pandion).
They are always horny (corneous). They take name from, and are
reckoned by, their respective digits: thus, 1 cl. = claw of U/ 2 cl.
- claw of 2t, etc.

Spurs (Lat. calcar, a spur) are developed on the metatarsal
bones of a few birds. They are of the nature of claws, being hard,
horny modifications of the epiderm : but they have nothing to do
with the digits. They possess a bony core upon which they are
supported, like the horns of cattle. Such growths chiefly occur in
gallinaceous birds : the spurs of the domestic fowl are a familiar
case. Sometimes there are a pair of such weapons on each foot, as
in the Pavo hicalcaratus, and there may be several more, as in the
genus Ithagenis. Another instance of their occurrence is offered
by the wild turkey (Meleagris gallipavo). Metatarsal spurs are
characteristic of the male sex they are offensive weapons, and
belong to the class of “ secondary sexual characters ” (p. 133). (For
wing-spurs, as shown in Fig. 53 ter, see p. 168.)

§ 4.— AN INTEODUCTION TO THE ANATOMY OF BIKDS

Anatomical Structure now affords ornithologists many and the
most important of the characters used in classification. In fact, few
if any of the groups above genera can be securely established without
consideration of internal parts and organs, as well as of exterior modi-
fications of structure. Therefore, the student who really “means
business ” must be on speaking terms at least with avian anatomy.
For example, none could in the least intelligently understand a
wing or a leg without knowing the bony framework of those
members. Yet to adequately set this matter forth would be to
occupy a very much larger volume with anatomy ; whereas, I can
onl}^ devote a few pages to the entire subject. In such embarrass-
ment, which attends any attempt to treat a great theme in a short
way that shall not also be a small way, attention must be mainly
confined to those points which bear most directly upon systematic
ornithology as distinguished from pure anatomy, in order to bring

GENERAL ORNITHOLOGY

PART II

forward the structures which are more particularly concerned in the
classification of birds. I wish to give a fair account of the skeleton,
as osteological characters are of the utmost importance for the deter-
mination of natural affinities ; and to continue with some notice of
prominent features of the muscular, vascular, respiratory, digestive,
urogenital, and nervous systems, and organs of the special senses,
as the eye and ear. The tegumentary system has already been
treated at some length ; so has the osseous system, so far as the
bones of the limbs are concerned. What further I shall have to
say is designed merely as an introduction to avian anatomy, and
is supposed to be addressed to beginners.

a. Osteology : The Osseous System, or Skeleton

Osteology (Or. oa-rkov, osteon, a bone ; X6yo^, logos, a word) is a
scientific description of bone in general and of bones in particular.
Bone consists of an animal basis or matrix (Lat. matrix, a mould)
hardened by deposit of earthy salts, chiefly phosphate of lime.
Bone is either preformed in the gristly substance called cartilage
(Lat. cartilago, gristle), and results from the substitution of the
peculiar osseous tissue for the cartilaginous tissue, or it is formed
directly in ordinary connective tissue, such as that of most mem-
branes or any ligaments of the body. Bone-tissue presents a peculiar
microscopic structure, in which it differs from teeth, as it does also
in not being developed from mucous membrane ; the substance is
called osteine, as distinguished from dentine. Though very dense
and hard, bone has a copious blood-supply, and is therefore very
vascular ; the nutrient fluid penetrates every part in a system of
vessels called Haversian canals. In the natural state bone is covered
with a tough membrane called periosteum (Gr. Trepl, peri, around, and
oa-rkov), which is to bone what bark is to a tree. The bones col-
lectively constitute the osseous system, otherwise known as the
skeleton (Gr. a-KeXerov, dried, as bones usually are when studied).
The skeleton is divided into the endoskeleton (Gr. evSov, endon, within),
consisting of the bones inside the body ; and the exoskeleton (Gr.
exo, out of), or those upon the surface of the body, of which birds
have none. Certain bones developed apart from the systematic
endoskeleton, in fibrous tissue, are called scleroskeletal (Gr. a-KXrjpos,
skleros, hard), as the ossified tendons or leaders of a turkey’s leg, the
ring of ossicles in a bird’s eye (an ossicle is any small bone).
Sesamoid (Gr. o-rja-apir], sesame, a kind of pea) bones, so often found
in the ligaments and tendons about joints, are also scleroskeletal.
The endoskeleton is divided into bones of the axial skeleton, so called
because they lie in the axis of the body, as those of the skull, back-

SEC. IV

ANATOMY OF BIRDS

199

bone, and chest ; and of the appendicular skeleton, including bones of
the limbs, considered as diverging appendages of the trunk. The
skeleton is jointed ; bones join either by immovable suture, or by
movable articulation (Lat. articulus, a joint, dimin. of artus, a limb).
In free articulations, the opposing surfaces are generally smooth,
and lubricated with a fluid called synovia. Progressive ossification
often causes bones originally distinct to coossify, that is, to fuse to-
gether ; this is termed ankylosis ; bones so melted together are said
to be ankylosed (Gr. dyKvXioa-is or d-yx^Awo-ts, the stiffening of joints
in a bent position). Thus all the bones of a bird’s brain -box are
ankylosed together, though this box at first consists of many
distinct ones ; and the determination of such osseous elements or
integers in compounded bones is a very important matter, as a clue
to their morphological composition. The names of most individual
bones, chiefly derived from the old anatomists, are arbitrary and
have little scientific signification ; many are fanciful and misleading ;
bones named since anatomy passed from the empiric stage, when it
was little more than the art of dissecting and describing, however,
have as a rule better naming. The shaft of a long bone is its con-
tinuity : the enlargements usually found at its extremities are called
condyles (Gr. kovSvAo?, kondulos, a lump, knot, as of the knuckles).
Points where ossification commences in cartilage or membrane are
ossific cenh'es, or osteoses ; valuable clues, usually, to the elements of
compound bones. But ossification of individual simple bones may
begin in more than one spot, and the several osteoses afterward grow
together. This is especially the case with the ends of bones, which
often make much progress in ossification before they unite with the
shaft or main part ; such caps of bone, as long as they are disunited,
are called epiphyses (Gr. €7rt, epi, upon ; pliusis, growth). Pro-

trusive parts of bones have the general name of processes, or apophyses
(Gr. diro, apo, away from, and covert?) ; such have generally no ossific
centres, being mere outgrowths. But many parts of a vertebra,
which are called “apophyses,” have independent ossific centres.
The progress of ossification is usually rapid and effectual.

The skeleton of birds is noted for the number and extent of its
ankyloses, a great tendency to coossification and condensation of
bone-tissue resulting from the energy of the vital activities in this
hot-blooded, quick-breathing class of creatures. Birds’ bones are
remarkably hard and compact. When growing, they are solid and
marrowy, but in after life more or fewer of them become hollow and
are filled with air. This pneumaticity (Gr. TrvevixarLKos, pneumatikos,
windy) is highly characteristic of the avian skeleton. Air penetrates
the skull-bones from the nose and ear-passages, and may permeate
all of them. It gains access to the bones of the trunk and limbs by
means of air-tubes and air-sacs, which connect with the air-passages

200

GENERAL ORNITHOLOGY

PART II

in the lungs ; such sacs, sometimes of great extent, are also found
in many places in the interior of the body, beneath the skin, etc.;
sometimes the whole subcutaneous tissue is pneumatic. The extent
to which the skeleton is aerated is very variable. In many birds
only the skull, in a few the entire skeleton, is in such condition ;
ordinarily the greater part of the skull, and the lesser part of the
trunk and limbs, is pneumatised. The passage of air in some cases
is so free, as into the arm-bone, for example, that a bird with the
windpipe stopped can breathe for an indefinite period through a

'Th'S

Fig. 54.— Ideal plan of the double-ringed
body of a vertebrate. neural canal ; if,
haemal canal ; the body separating them is
the centrum of any vertebra, bearing e, an
epapophysis, and y, a hypapophysis ; n, n,
neurapophyses ; d, d, diapophyses ; ns,
bifid neural spine ; pi, pi, pleurapophyses ;
h h, hsemapophyses ; hs, bifid haemal spine.
Drawn by Dr. R. W. Shufeldt, U.S.A., after
Owen.

Fig. 55. — Actual section of the body in the
thoracic region of a bird. N, neural canal ; if,
haemal canal ; c, centrum of a dorsal vertebra ;
hy, hypapophysis ; d, diapophysis ; z, zygapo-
physis ; ns, neural spine ; r, pleurapophysis, or
vertebral part of a free rib, bearing u, uncinate
process or epipleura ; cr, haemapophysis or
sternal part of the same ; st, section of sternum
or breast-bone (haemal spine). Designed by Dr.
R'. W. Shufeldt, U.S.A.

hole in the humerus. Pneumaticity is not directly nor necessarily
related to power of flight ; some birds which do not fly at all are
more pneumatic than some of the most buoyant. (On the general
pneumaticity of the body see beyond, under head of the Kespiratory
System.)

The Axial Skeleton (Figs. 54, 55, 56) of a bird or any vertebrated
animal, that is, one having a back-bone, exhibits in cross-section two
rings or hoops, one above and the other below a central point, like
the upper and lower loops of a figure 8. The upper ring is the
neural arch (Gr. vevpov, neuron, a nerve), so called because such a

SEC. IV

ANATOMY OF BIRDS

201

cylinder encloses a section of the cerebro-spinal axis, or principal
nervous system of a vertebrate (brain and spinal cord, whence arise

all the nerves of the body, excepting those of the sympathetic
nervous system). The lower ring is the Ticemal arch (Gr. atfia, haima,

202

GENERAL ORNITHOLOGY

PART II

blood), which similarly contains a section of the principal blood-
vessels and viscera. Fig. 55 shows such a section, made across the
thoracic or chest region of the trunk. Here the upper ring (neural)
is contracted, only surrounding the slender spinal cord, while the
lower ring is expanded to enclose the heart and lungs. Such a
section, made in the region of the skull, would show the reverse ;
the upper ring greatly inflated to contain the brain, the lower con-
tracted and otherwise greatly modified into bones of the jaws. Thus
the trunk of a vertebrate is a double-barrelled tube ; one tube
above for the axial nervous system, the other below for the viscera
at large ; the partition between the two being a jointed chain of
solid bones from one end of the body to the other. These solid
bones are the centrums or bodies of vertebrce, in the trunk ; and in the
head certain bones which in some respects correspond with the
centrums of vertebrae. The entire chain or series of vertebrae com-
poses the back-bone or spmal column ; with its connections (thorax)
and anterior continuation (skull) it is the axial skeleton. The skull
is considered by some competent anatomists to consist of modified
vertebrae. The skull-bones have certainly the position and relations
of parts of vertebrae ; to a certain extent they resemble vertebrae,
as in being divisible into several segments, like as many vertebral
segments ; they are also directly in the axis of the body, enclosing
a part of the cerebro-spinal nervous system above, and portions of
the visceral systems below. But supposed strict morphological
correspondence of cranial bones with vertebrae is not supported by
their mode of development, and is now generally denied, the relation
being considered rather analogical and physiological than homolo-
gical and morphological.

1. THE SPINAL COLUMN

A Vertebpa (so called from the flexibility of the chain of
vertebrae ; Lat. verto^ I turn) consists of a solid body or centrum,
and more or fewer processes or apophyses, some of which have
separate ossific centres. Plate-like processes which arch upward
from either side of a centrum to enclose the neural canal are the
neural arches or neurapophyses (Fig. 54, n, n) ; at their union in the
middle line above they commonly send up a process called the
neural spine (ns). Transverse processes from the sides of the neural
arch are diapophyses (Gr. §ta, dia, across ; Figs. 54, 55, d, d). Oblique
processes from the sides of the same arches, serving to lock them
together, are zygapophyses (Gr. fuyov, zugon, a yoke ; Fig. 55, z)-,
there are two on each side ; one anterior, on the front border of an
arch, a prezygapophysis ; one posterior, on the hind border, a post-
zygapophysis. From the under side of a centrum, in the middle

SEC. IV

ANATOMY OF BIRDS

203

line, there is often a liypapo]phjsis (Gr. viro, hupo, under ; Fig. 55, liy).
These several processes, with some others not necessary to mention
here, make with the centrum a vertebra in strictness ; that is, when
existing at all, they are completely consolidated with one another
and with the centrum into one bone. But certain important ele-
ments of a vertebra, developed from independent ossific centres,
may or may not ankylose therewith, in different regions of the
same spinal column. These are the pleurapopliyses (Gr. TvXevpov,
pleuron, a rib; Fig. 54, pi ; Fig. 55, r). Airy rib is in fact the pleura-
pophysial element of a vertebra ; it may be, and in most regions of
the spinal column it is, quite small when existing at all, and
ankylosed with the vertebra to which it belongs, as an integral
portion thereof. But in the lower region of the neck, and through-
out the thoracic region, such pleurapophyses elongate, and are
movably articulated with their respective vertebrae ; they then
become the “ribs” of ordinary language. Moreover, the true
thoracic ribs of birds are jointed near the middle, each thus con-
sisting of two pieces ; the upper piece is pleurapophysis proper :
the lower is called a hcemapopliysis (Fig. 54, h ; Fig. 55, cr); it corre-
sponds to a “ costal cartilage ” of human anatomy. Once again :
since the sternum (breast -bone) is theoretically, and doubtless
archetypically, a solidified set of those parts of the vertebral
segments which complete the haemal arches below, each segment of
a sternum to which a haemapophysis is articulated is called a Immal
spine, being compared to a neural spine above. Aside from any
consideration of the ribs proper and sternum, or free pleurapophyses,
haemapophyses, and haemal spines, any “vertebra” of ordinary
language is the compound bone which consists of centrum and
neur-, di-, pre-, and post-zyg-, pleur-, hyp-, and other -apophyses,
if any, and neural spine; the latter often called “spinous
process.”

The Vertebrae join one another, forming a continuous chain.
Their centra are placed end to end, one after another ; their neural
arches are also locked together by the zygapophyses, when these
articular processes are developed. Zygapophyses bear upon their
free ends smooth articular facets, the faces of which are mostly
horizontal ; those of the prezygapophyses looking downward, and
overriding the reversed faces of the postzygapophyses. The mode of
jointing of the centra of such vertebrae as are freely movable upon
each other is highly characteristic of birds, in so far as the shapes of
the articular ends of the vertebral centra are concerned. In anatomy
at large, a vertebral centrum which is cupped or hollowed at both
ends is of course biconcave. Such a vertebra is called amphicoelous
(Gr. dfjL(f>L, amphi, on both sides ; kolXos, koilos, hollowed) ; this is
the rule in fishes, and obtained in some extinct Cretaceous birds, as

204

GENERAL ORNITHOLOGY

PART II

Ichthyornis ; it is unknown in recent birds. ^ A centrum cupped in

front only is procodous ; one cupped only behind is opisthocodous
(Gr. oTTLo-Oe, opisthe, behind). Such structure results in a ball-and-
socket jointing of vertebrae. In those vertebrae of birds in which
this arrangement obtains, it is always the posterior face of a centrum
which is cupped, the anterior one being balled ; such vertebrae are
therefore opisthocoelous. But in the freest vertebral articulation of
birds, that existing in the region of the neck, another modification
occurs. Both ends of each vertebra are saddle-shaped, i.e. concave
in one direction, convex in the other ; a condition which is called
heterocoelous (Gr. eVepos, lieteros, contrary). The concavo-convexity of
any one vertebra fits the reciprocal concavo-convexity of the next.
Anterior faces of heterocoelous vertebrae are concave crosswise, up-
and-down convex ; posterior faces are the reverse : consequently,
such vertebrae are procoelous in horizontal section, but in vertical
section opisthocoelous. The various physical characters of vertebrae
in different regions of the body, and their connections with and
relations to other parts of the body, have caused their division into
several sets, as cervical, dorsal, etc., which are best considered
separately.

Cervical Vertebrae (Fig. 56, cv) are those of the neck: all those
in front of the thorax or chest, which do not bear free pleura-
pophyses in adult life, or the free pleurapophyses of which, if any,
are not in two- jointed pieces and do not reach the breast-bone ; i.e.
have no hsemapophyses. It is advisable, in birds, to draw this line
between cervical and succeeding vertebrae, no other being equally
practicable ; for, on the one hand, one, two, or more of the cervicals
(recognisable as such by their general conformation and free articu-
lation) may have long free ribs, movably articulated ; and all the
cervicals, excepting usually the first, or first and second, have short
pleurapophyses, ankylosed in adult life, but free in the embryo ;
while, on the other hand, a vertebra, apparently dorsal by its con-
figuration and even its ankylosis with the dorsal series, may be
entirely cervical in its pleurapophysial character.^ Thus, in Fig. 56,
of an owl’s trunk, the bone which is apparently first dorsal, and is
so marked {dv\ bears a free styliform “ riblet ” an inch long (c'), only

^ Except to this statement, however, the oddly-massed pygostyle, which, in birds
where a terminal disk develops inferiorly, may be distinctly cnj^ped at both ends, as
it is in a raven, for example.

2 The case is very puzzling ; the more so because, viewing the whole series of
birds, the ambiguous “ cervico-dorsal,” or two such equivocal vertebrae, may lean in
different cases in opposite directions when the whole sum of characters is taken into
account. Therefore it may be best, as already said, to make the possession of a
jointed sternum-reaching rib the criterion of the first dorsal vertebra, even though an
antecedent one may have the physical characters of a dorsal, and be ankylosed with
the dorsal series. This is the view taken by Huxley, who says ; “ The first dorsal
vertebra is defined as such by the union of its ribs with the sternum by means of a

SEC. IV

ANATOMY OF BIRDS

205

it is not jointed, and does not reach the sternum ; while the next
to the last cervical has a minute but still free rib (c). In a raven’s
neck before me, the last cervical rib is about two inches long,
articulating by well-defined head and shoulder to body and lateral
process of the vertebra ; the penultimate rib is about half an inch
long, with one articulation to the lateral process ; while the next
anterior vertebra (third from the last) has a minute ossicle, as a free
“riblet.” The rule is two such free pleurapophyses or cervical ribs
of any considerable length : sometimes one ; rarely, three ; in the
cassowary four. Eudimentary pleurapophyses may usually be
traced up to the second cervical vertebra, as slender stylets or
riblets, completely ankylosed with the neural arches in adult life,
and lying parallel with the long axes of the bones. The ankylosis
of pleuropophyses distinguishes most cervical vertebrae in another
way : for from it results, on each side of the neural arch, a foramen
(Lat. foramen, a hole, pi. foramina), through which blood-vessels
(vertebral artery and vein) pass to and from the skull. The series
of these foramina is called the vertebrarterial canal ; none such exist
in those posterior cervical vertebrae which bear free ribs j thus, in
the raven the canal begins abruptly at the fourth from the last
cervical. But, as in Rhea, for instance (and doubtless in many
other cases), the vertebrarterial canal shades visibly into the series
of foramina formed by the spaces between the head and shoulder
of any rib and the side of the vertebra to which it is attached ;
such being the true morphology of the canal. The cervical is the
most flexible region of a bird’s spine ; the articular ends of the
vertebral bodies are the most completely saddle-shaped (hetero-
coelous) ; the zygapophyses are large and flaring, overriding each
other extensively ; the largest processes are at the fore ends of the
bones ; the appositions of the central and zygapophysial articular
surfaces are collectively such, that the column tends to bend in an
S-shape or sigmoid curve. The vertebral bodies are more or less
contracted in the middle, or somewhat hourglass-shaped ; on several
lower cervicals hypapophyses are likely to be well developed ; as
are neural spines toward both the beginning and end of the series.
The vertebrae on the whole are large ; their neural canal is also of
ample calibre. The first two cervicals are so peculiarly modified
for the articulation of the skull as to have received special names.

sternal rib.” {Anat. Vert. Anim., 1872, p. 237.) Owen appears to regard as dorsal
any of the vertebrae in question which bear free ribs. The actual uncertainty in the
case, and the discrepant reckoning by different authors, prevents us from making
a satisfactory count of the numbers of the two series of vertebrae in any given case.
Thus, Fig. 56, as marked by Dr. Shufeldt, shows six dorsals {dv), to which is to be
added the one under p, bearing the rib sr ; and from which is to be subtracted the
anterior one, bearing the rib c\ which is to be regarded as cervical, though its
physical characters are evidently those of the dorsal series.

2o6

GENERAL ORNITHOLOGY

PART II

The first one, Fig. 56, at^ the atlas (so called because it bears up the
head, as the giant Atlas was fabled to support the firmament), is a
simple ring, apparently without a centrum. The lower part of the
ring is deeply cupped to receive the condyle of the occiput into a
ball-and-socket joint. The second cervical is the axis, ax, which
subserves rotary movements of the skull. It has a peculiar tooth-
like odontoid (Gr. 68ov<s, oSovto?, odous, odontos, tooth ; e?Sos, eidos,
form) process, borne upon the anterior end of its body, fitting into
the lower part of the atlantal ring ; about which pivot the atlas,
bearing the head, revolves like a wheel upon an eccentric axis.
The cervicals of birds vary greatly in number ; according to Huxley
there are never fewer than eight, and there may be as many as
twenty- three ; Stejneger gives twenty-four for some of the swans.
Twelve to fourteen may be about an average number.

Thoracic or Dorsal Vertebrae (Fig. 56, dv) extend from the
cervical to or into the pelvic region of the spine. In most animals,
and in ordinary anatomical language, a “ dorsal ” is one which bears
a distinct free rib, and is therefore truly thoracic, since “ ribs ” are
the side- walls of the chest. But in birds, as we have seen, certain
cervicals have distinct elongate ribs ; and, as will be seen soon, long
jointed pleurapophyses are usually found in that region commonly
called “sacral.” The first dorsal, in birds, is arbitrarily considered
to be that one which bears the first rib which is jointed, and which
reaches the sternum by its lower (hsemapophysial) half. Five or
six vertebrae of birds commonly answer this description ; though
the last one which bears a long free jointed rib (which may or may
not reach the sternum) is commonly ankylosed with the sacrum, as
sr. So few as only three haemapophysis-bearing ribs may reach the
sternum. There may also be a long free-jointed rib which “ floats ”
at loth ends ; i.e. is articulated neither with the sternum nor with
the vertebra to which it belongs, as in the loon, for example. As
the dorsal series thus shades insensibly behind into another series,
the lumbar (which has no free, nor any distinct ribs, — ribs that one
would not hesitate to call such), it is best to consider as dorsal or
thoracic all those vertebrae, succeeding the last cervical (which is to
be determined as explained in the last paragraph), which have
distinct jointed ribs, whatever the connection or disconnection of such
pleurapophyses at either end. On this understanding, one, some-
times two or even three “ dorsal ” vertebrae ankylose with the pelvic
region of the spine. Fixity of the dorsal region being of advantage
to flight, these vertebrae are very tightly locked together ; not only
by the close apposition or even ankylosis of their bodies and pro-
cesses, but also, in many cases, by ossifications of the tendons of
muscles of the back, and coossifications of these with the vertebrae,
like a set of splints, till the consolidation of the thoracic is only

SEC. IV

ANATOMY OF BIRDS

207

surpassed by that of the pelvic region of the spine. Dorsal
vertebrae also usually differ a good deal from most cervicals in
having shorter bodies, laterally compressed, producing a ridge
which runs along their middle line below ; in lacking a vertebrar-
terial canal ; in having on each side two articular facets, — one on
the body and the other on the transverse process, for the head and
shoulder of a rib. They are further distinguished, usually, by
having large spinous processes, in the form of high, long, thin,
squarish plates, often or usually ankylosed together. Their trans-
verse processes are also very prominent laterally, thin and
horizontal, and often ankylosed. More or fewer dorsals may bear
large hypapophyses ; which, as in the loon, may bifurcate at their
ends into two flaring plates. Such processes continue a similar
series from the neck, and are in relation to the advantageous action
of the muscles {rectus colli anticus and longus colli) by which the neck
is made to straighten out from the lower curve of its sigmoid
flexure.

The “ Sacrum ” of a Bird (Figs. 57 and 60) is commonly con-
sidered to be that large solid mass of numerous ankylosed vertebrae
in the region of the pelvis, covered in by, and fused more or less
completely with, the principal bones of the pelvis, or haunch-bones
{ilia). But in this consolidation of an extremely variable number
(averaging perhaps twelve, but running up to at least twenty, eleven
to thirteen being usual) of • bones are included vertebrae which in
other animals belong to several different sets — dorsal, lumbar,
sacral proper, and coccygeal or caudal. We have just seen that one
or two, even three, vertebrae, which are dorsal according to the
definition agreed upon, may enter into the composition of the
“sacrum,” being firmly ankylosed therewith, and their long ribs
issuing out from underneath the ilia, as shown in Fig. 56, sr. Next
comes one bone, or a series of several (two to five or more) bones,
ankylosed together by their bodies and spinous processes, and also
ankylosed with the ilia by means of stout lateral bars of bone sent
transversely outward on either side from their respective centra to
abut against the ilia. These cross-bars correspond in general form
and position with the transverse process of the last true rib-bearing
dorsal, — that process against which the shoulder of any developed
rib abuts ; they are variously considered to be, to represent, or to
include, rudimentary ribs j and such difference of view may be war-
ranted by the state of the parts in different birds. However this
may be, the bones just described are lumbar vertebrae (Lat. lumbus,
the loin; where such vertebrae are situated in man and other
mammals) ; which certainly possess abortive ribs in some cases.
On successive lumbars the cross-bars, whatever their nature, com-
monly slip lower and lower downward (belly-ward) on the vertebral

208

GENERAL ORNITHOLOGY

PART II

bodies, till the last ones are quite down to the level of the ventral
aspect of the centrum these are also commonly the stoutest, most
directly transverse, and most nearly horizontal of the series of pro-
cesses, abutting against the ilia a little in advance of the socket of
the thigh-bone. This ends a series of consolidated “ sacral ” verte-
brae which are termed collectively “ dorso-
lumbar,” — all of them anterior to the
true sacrum of a bird. The sacrum ^proper
(Fig. 57, s) consists of those few vertebrae
— three, four, or five — from foramina be-
tween which issue the spinal nerves that
form the network called the sacral plexus.
These true sacral vertebrae are ribless, and
may be recognised, in a general way, by
the absence of anything like the cross-bars
above described, issuing from the verte-
bral centra ; though their neural arches
send off some small bars or plates to fuse
with the ilia. These sacrals proper are at
or near the middle of the whole sacral
mass. After these come a large number
— from five to ten or more — of vertebrse
which, from their following the true
sacrals, though consolidated therewith
and with one another, are considered to
belong to what would be the caudal
region of other animals, and are hence
called tail-sacrals,” urosacrals (Gr. ovpa,
tail. Fig. 57, c). These continue to send
off a series of little plate-like processes
from their neural arches, just as the true
sacrals do ; but, in addition to these, pro-
cesses are given off from the bodies of
the urosacrals, corresponding in position
and relation to those which proceed from
the bodies of the lumbars, and being ap-
parently of the same morphological char-
acter (pleurapophysial). These “ riblets ”
are, however, quite slender, and also
oblique in two directions j for instead of being transverse and
nearly horizontal, they trend very obliquely backward and
upward j they also shorten consecutively from before back-
ward. The cross - bars of the latter urosacrals, however, are
stouter and altogether more like those of a lumbar vertebra.
The appearances described are those seen from below, or on

Fig. 57. — The sacrarium of a
young fowl, seen from below, nat.
size; after Parker, dl, ilorsolum-
bar series, whereof the first is dor-
sal proper, the next three are lum-
bar ; s, the sacral series proper, or
true sacrum, consisting of five
vertebrse ; c, the urosacral series,
being those caudal vertebrae, six in
number, which anchylose with one
another and with the sacrum.

SEC. IV

ANATOMY OF BIRDS

209

the ventral aspect. Above, on the back of the pelvis, the line of
confluent spinous processes of the dorsolumbars is commonly dis-
tinct, separated a little from the flaring lips of the ilia. Such dis-
tinct formation may continue throughout the sacral and urosacral
regions ; oftener, however, the line of spinous process sinks, flattens,
and widens into a horizontal plate which becomes perfectly con-
fluent with the ilia along the posterior portion of their extent ; such
smooth, somewhat lozenge-shaped surface being quite continuous
with the superficies of the pelvis, but perforated with more or fewer
pairs of intervertebral foramina. — Such is the general character of a
bird’s complex sacrarium, as I name the whole mass of bones that are
ankylosed together, including dorsolumbars and urosacrals, as well
as sacrum proper. The description is taken chiefly from a raven
(Corvus corax) ; the figure from the common fowl, after Parker.
The kidneys are moulded into the recesses between the sacral and
urosacral vertebrae and in the concavity of the ilia. The general
shape of a sacrarium, viewed from below, is fusiform, broadest
across the sacral bodies proper or just in front of them, tapering to-
ward either end ; the face of the sacrarium is also flattest about the
middle, more or less ridged before and behind from compression of
the vertebral bodies. It has little if any lengthwise curvature, and
that chiefly in the urosacral region, where the concavity is down-
ward. The total number of bones may be less than twelve, or more
than twenty. The extensive ankyloses in this region of the spine
are in evident adaptation to bipedal locomotion, which requires
fixity hereabouts, that the trunk may not bend upon the fulcrum
represented by a line drawn through the hip-joints, which are
situated about opposite the middle of the sacral mass, as shown by
the arrow, ac, in Fig. 60.

The Coccygeal, or Caudal Vertebrse (Fig. 56, civ) proper, ter-
minate the spinal column. They are called “ coccygeal,” from the
fancied resemblance of the human tail-bones collectively to the beak
of a cuckoo (Gr. kokkv^, JcoJckux). The caudals are all the free bones
situated behind the ankylosed urosacrals. The series commonly
begins opposite the point where the pelvic bones end ; it consists of
a variable number of bones, from the twenty long slender ones
which the Archaeopteryx possessed, down to seven or fewer separate
ones. The usual number is eight without the pygostyle. They are
stunted, degraded vertebrse, whose chief office is to support the tail-
feathers : for the leash of nerves which emerge from the spinal
canal to form the sacral plexus by so much diminishes the spinal
cord that a mere thread is left to penetrate the tail, though the
neural arches of all the coccygeals be still pervious. All may be
freely movable, as in the American Ostrich {Rhea) ; but in almost
all birds only the anterior ones are distinct and vertebra-like, the
p

210

GENERAL ORNITHOLOGY

PART II

rest, to a variable number, being abortive, and melted into that
extraordinary affair called the rump-post or ;pygostyle (Gr. irvyr^,
the rump ; o-rvXog, a post), which may consist of no fewer than
ten such metamorphosed tail-bones. It has usually a shape sug-
gesting the share of a plough (see Fig. 56, ^y), but is too variable
to be concisely described. The pygostyle supports the tail-feathers ;
and as these are morphologically one pair to each rectrix-bearing
vertebra, the number of tail-feathers may be primarily equal to the
number of vertebrae which fuse in the pygostyle. Thus the swan
is said to have ten vertebrae in this mass ; a wild swan {Cygms
columbianus) has twenty tail-feathers. In this view, six should be
the usual composition of the share-bone. A bird’s tail is really
more extensive and lizard-like than commonly supposed ; thus the
swan, besides its ten in the pygostyle, has seven free caudals, and
ten urosacrals — twenty-seven post-sacral vertebrae in all (Huxley).
In the raven, the free caudals are six, exclusive of the pygostyle.
These all have large flaring transverse processes and moderate
spinous processes, and the latter ones are also provided with hypa-
pophyses, some of which are bifurcate. The pygostyle in many
birds expands below into a large circular or polygonal disk.

2. THE THORAX: RIBS AND STERNUM

The Thorax (Gr. Ocopa^, a coat of mail ; in anatomy, the chest ;
adj. thoracic ; see Fig. 56) is the bony box formed by the ribs on each
side, the breast-bone below, and the back-bone above. In birds it
is very extensive, including most or all of the abdominal as well as
the thoracic viscera, and its cavity is not partitioned off from that
of the belly by a completed diaphragm^ though a rudimentary struc-
ture of that kind is found in the Apteryx. The thorax is usually
soldered behind to the pelvis by union of one or more pairs of ribs
with the ilia ; in front it always and entirely bears the pectoral arch
(see p. 215). The thorax is very movable in birds, by reason of
the great length and jointedness of the ribs.

The Ribs (Lat. costa, a rib; pi. costoe; adj. costal; see Fig. 56,
c, c', R, cr, sr, u), as said above, are the pleurapophysial elements of
vertebrae, which remain small and ankylosed, or become long and
free. In the latter state only are they “ ribs ” in ordinary language.
The one or more cervical ribs, however elongated, and the abortive
lumbar and urosacral ribs, are to be excluded from the present
description, and have been already considered. True ribs are those
which belong to the dorsal vertebrae proper, and are jointed in
themselves ; that is, have articulated hcemapophyses (see p. 203), by
which they may or do articulate with the sternum. Such true ribs
are fixed, when they reach from back-bone to breast-bone ; fioating.

SEC. IV

ANATOMY OF BIRDS

2II

when either or neither of these connections is made. Usually the
last rib, though hearing a perfect hsemapophysis, does not reach the
sternum ; in the loon, for example, the last rib floats at both ends,
having connection neither with vertebra nor sternum ; and the two
next ribs float at their sternal ends. The perfected ribs are few, —
flve or six is a usual number, though nine are hsemapophysis-bearing
in the loon. The last rib at least is usually sacrarial, as it belongs
to a dorsal vertebra which is ankylosed with the sacrarial mass ;
and two, or even, as in the loon, three ribs may likewise issue out
from under cover of the ilia. These “sacral ribs” or sacrocostals are
furthermore distinguished by being devoid of the epipleural or un-
cinate ^processes (Lat. uncus^ a hook ; Fig. 5 6, u) with which other true
ribs are furnished, forming a series of splint-bones proceeding
obliquely from one rib to shingle over the next succeeding one,
and thus increase the stability of the thoracic side-walls. Such
splints may be either articulated or ankylosed with their respective
ribs ; they have independent ossific centres. The upper (pleura-
pophysial) part of a rib, or “ vertebral rib,” when perfected, articu-
lates with the side of the body of a vertebra by its head or capit-
ulum (Lat. dimin. of caput, head), and also with the lateral process
of the same vertebra by its shoulder or tuberculum (Lat. dimin. of
tuber, a swelling). In well-marked cases the head and shoulder are
quite far apart, the rib seeming prolonged above ; either of these
vertebral connections may be disestablished, the other remaining, or
both may be lost. The lower (hsemapophysial) part of a rib, or
“ sternal rib,” articulates with the side of the sternum by a simple
enlargement ; the ends of those sternal ribs which thus join the
sternum tend to cluster closely together at a part of the breast-bone
called its costal process (Fig. 58) ; those which do not make the
sternal connection are simply bundled together. Commonly five or
six, sometimes four, rarely only three ribs reach the sternum. The
ribs are ordinarily as slender and strict as those shown in Fig. 56 ;
but in Apteryx, for example, their pleurapophysial parts are expan-
sive and plate-like. They lengthen rapidly from before backward,,
both in their vertebral and their sternal moieties ; these parts meet
at angles of decreasing acuteness from before backward ; but these'
angles, as those of the ribs both with vertebrae and sternum, inces-
santly increase and diminish in the respiratory movements of tho
chest j all being in expiration more acute, and more obtuse in
inspiration.

The Avian Sternum (Gr. cnepvov, sternon,thQ breast; Fig. 56, /S')
is highly specialised ; its extensive development is peculiar to the
class of birds, and its modifications are of more importance in
classification than those of any other single bone. Thereupon it
becomes an interesting object. Theoretically it is a collection of

212

GENERAL ORNITHOLOGY

PART II

haemal spines of vertebrae. Though such morphological character
is appreciable in those animals which have a long jointed sternum,
the segments of which, answering to pairs of ribs, develop from
separate centres, there is little or nothing in the development or
physical characters of the avian sternum to favour this view. The
great bone floors the chest and more or less of the belly, and fur-
nishes the main dJa'p^ui of both the bony and muscular

apparatus of flight, receiving important bones of the scapular arch
and giving origin to the immense pectoral muscles. (See also Fig.
58.)

Birds offer two leading types of sternal structure, the mtite and
the carinate, or the “raft-like” and the “boat-like,” according as
the bone is flat or keeled (Lat. ratis, a raft ; adj. ratite ; in an arbi-
trary nom. pL, Ratitce, a name of one of the leading divisions of
birds : Lat. carina, a keel ; adj. carinate : nom. pi. Carinatce, name
of another such division). 1. In all struthious birds, comprehend-
ing the ostrich and its allies (and also in the Cretaceous Hesperornis),
the sternum is a flattish, or rather concavo-convex, buckler-like bone,
of somewhat squarish or rhomboidal shape, developed from a single
pair of lateral centres of ossification — a “flat-boat,” without any
keel, built with reference to an important modification of the
shoulder-girdle, and a reduced or rudimentary condition of the
wings, which are unfit for flight. 2. In all flying birds, and some
which from other than any fault of the sternum do not fly — com-
prising all remaining recent birds, or Carinatce, and also the Creta-
ceous Idithyornis — the sternum is keeled and develops from a
median centre of ossification as well as from lateral paired centres ;
usually two of these, making five in all. In a few CarinatcE the
keel is rudimentary, as the flightless ground parrot of New Zealand,
Stringops habroptilus ; or otherwise anomalous, as in the extra-
ordinary Opisthocomus cristatus, where it is cut away in front, and
in the rail-like Notornis, where the sternum is extremely like a
lizard’s. In general, the development of the keel is an index of
wing-power, whether for flying or swimming, or both ; the effective-
ness of the pectoral muscles being rather in proportion to depth
of keel than to extent of the sides of the “ boat-bone ” j thus,
the keel is enormous in swifts (Cypselidce) and humming-birds
{Trochilidm).

The carinate sternum normally develops from five centres,
having consequently as many separate pieces in early life. Two of
these are lateral and in pairs j the third is median and single. The
median ossification, which includes the keel, is the lophosteon (Cr.
Ao<^os, lophos, a crest ; ocrreov, osteon, a bone). The anterior lateral
piece, that with which the ribs, or some of them, articulate, is the
pelurosteon (Gr. TrXevpbv, pleuron, a rib) ; in adult life this becomes

SEC. IV

ANATOMY OF BIRDS

213

the costal process, so prominent in Passeres (Fig. 58). The posterior
lateral piece is the metosteon (Gr. [xera, meta, after). From the
latter are derived the pair, or two pairs, of lateral processes which
the posterior border of the sternum has in many birds. In fine, the
extent of ossification of the lophosteon and metostea, and the mode
of their coossification, determines all those various shapes of the
posterior border of the sternum which, being commonly character-
istic of genera and higher groups, are described for purposes of
classification. Thus, if the lophosteon and the metostea are com-
pletely ossified and to the same extent behind, the posterior border
of the sternum will be transverse, and perfectly bony. Such a
sternum is said to be entire. If the lophosteon is longer than
the lateral pieces, the sternum will have a median pointed or
rounded projection ; such a formation is called the middle xiphoid
process (Gr. xiphos, a sword : elSos, eidos, form). The pro-

jection of the metostea, not infrequent, similarly gives a pair of
external lateral xiphoid processes. But such processes oftener
result merely from defects of coossification between the elements of
the sternum. Thus, there is often a deep notch in the posterior
border of the sternum between the lophosteon and the metosteon
of each side ; the sternum is then said to be single-notched or single-
emarginate (one pair of notches, one on each side; Fig. 58). This
conformation prevails throughout the great group Passeres, possibly
without exception ; it is therefore highly characteristic of that
order, though a great many other birds also have it. In the
natural state, the notch is filled in with membrane. Such a notch
may also be converted into a “ fontanelle ” or fenestra (Lat. fenestra,
a window), which is simply a hole in the bone, the metostea having
grown to the lophosteon at their extremities, but left an opening
between. Such a sternum is called fenestrate, more exactly uni-
fenestrate (Lat. unus, one ; one window on each side). Now, the
parts remaining as before, let each half of the lophosteon, or each
metosteon, be notched or fenestrate ; obviously, then, such a sternum
is double-notched or bi-fenestrate, having four notches, or holes, two on
each side — two notches, or two holes ; or notched and fenestrate,
having a notch and a hole on each side. The latter is very fre-
quent ; when occurring, the hole is generally nearest the middle
line, the notch exterior. Irregularity of ossification, converting a
hole into a notch, and conversely, may in any case result in lack of
symmetry ; but this is a mere individual peculiarity. When there
are two notches on each side, as in Fig. 56, the sternum has evi-
dently a median and two lateral backward extensions, which are
then called respectively the middle, internal lateral, and external
lateral xiphoid processes. Notching of the lophosteon in the middle
line, at least to any extent, must be very rare, if indeed it ever

214

GENERAL ORNITHOLOGY

PART II

occurs. The extreme case of emarginatioii of the sternum is afforded
by the Gallince, and is highly characteristic of that group. Here
the lophosteon is extremely narrow, and fissured deeply away from
the metostea, which latter are deeply forked ; the arrangement
giving rise to two very long slender lateral processes on each side
(Figs. 1 and 2, p. 73). The sternum of the tinamou, a dromseo-
gnathous bird, is still more deeply emarginated, but the extremely
long and slender lateral processes, which enclose an oval contour,
are simple, not forked.

In a very few birds there are centres of ossification additional
to those above described. In Turnix, there are said by Parker to

be a pair of centres between the pleur-
ostea, which he names coracostea, because
related to the part of the sternum with
which the coracoids (see p. 216) unite.
The same authority describes for Dicho-
lophus a posterior median cartilaginous
flap having a separate centre, named
^ ur osteon (Gr. oppa, our a, tail). In various
birds the sternum is eked out in the
middle line behind by cartilage which
has no ossification.

Fig. 58.— Typical passerine Well marked in Passerme birds (Fig. 58) :

sternum, pectoral arches, and ster- ■, t '

nai ends of ribs ; from the robin, and may be bif urcate at the end and run

R’’wf down the front of the keel some way, as

single - notched, with prominent in the raven. The fore border of the

nnafnl nrnppsHPs anfl frirlrprl mnnii.

have prominent pleurostea, produced in angular costal processes. This
border is also thickened,' and presents on each side a well-marked,
smooth-faced groove, in which the expanded feet of the coracoid bones
are instepped and firmly articulated. These deep grooves commonly
meet in the middle ; are occasionally continuous from one side to
the other ; sometimes each crosses to the other side a little way.
The costal processes on each side also have thickened edges, with
a series of articular facets for the ribs, which gives this border a
fluted or serrate profile. Generally the fore half, or rather less, of
the side border of the sternum is thus articular ; and it is only such
costiferous (rib-bearing) extent of sternum which corresponds to the
whole body of the bone in a mammal, all the rest being “xiphoid.”

The sternum, especially of the higher
birds, develops in the middle line in
front a beak - like process called the
rostrum or manubrium (Lat. manubrium^
a handle) ; its size and shape vary ; it is

side to side, and then, in those birds which

SEC. IV

ANATOMY OF BIRDS

215

The singular carinate sternum of Notornis, and the ratite hone of
Ajpteryx, are concave crosswise along the front border, and bear the
coracoids far apart, at the summits of antero-lateral projections.

A sternum is generally concavo-convex in every direction,
bellying downward ; somewhat rectangular, it may be long and
narrow, or short, broad, and squarish. It is commonly longer than
broad, with convex front border, a median beak, which is often
forked, prominent antero-lateral corners, pinched-in sides (bulging
in tinamou) and indeterminate hind border. The keel usually
drops down lowest in front, sloping or curving gently up to the
general level behind, with a concave (rarely protuberant) vertical
border, and pronounced apex, to which the clavicles may or may
not he ankylosed, as they are in a pelican, for instance. In Opis-
thocomus the clavicles ankylose with the manubrium of the sternum.
The external surface, both of body and keel, is ridged in places,
indicating lines of attachment of the different pectoral muscles.
In a few birds, notably swans and cranes, the sternal keel is ex-
panded and hollowed out to receive folds of the windpipe in its
interior (see Figs. 99, 100). — But the numberless modifications of
the sternum in details of configuration belong to systematic orni-
thology, not to rudimentary anatomy.

3. THE PECTORAL ARCH

The Peetopal Apch (Lat. pectus, the breast; Figs. 1, 2, 56, 58,
59) is that bony structure by which the wings are borne upon the
axial skeleton. It is to the fore limb what the pelvic arch is to the
hind limb ; but is disconnected from the back-bone and united with
the breast-bone, whereas the reverse arrangement obtains in the
pelvic, which is fused with the sacral region of the spine. Each
pectoral arch of birds consists (chiefly) of three bones : the scapula
and coracoid, forming the shoulder-girdle proper, or scapular arch ; and
the accessory clavicles, or right and left half of the clavicular arch.
There is also at the shoulder-joint of most birds an insignificant
sesamoid ossicle, called scapula accessoria or os humero-scapulare (Fig.
56, ohs)', and in many a rudiment of a bone called procoracoid,
which occurs in reptiles, but in birds is united with the clavicle.
From the ribs, the scapula; from the sternum, the coracoid; from
its fellow, the clavicle, converges to meet each of the two other
bones at the point of the shoulder. The lengthwise scapular arches
of opposite sides are distinct from each other ; the clavicular arch is
crosswise, and nearly always completed on the middle line of the
body ; by which union of the clavicles the whole pectoral arch is
coaptated. The coracoid bears the shoulder firmly away from the
breast ; the scapula steadies the shoulder against the ribs ; the

2I6

GENERAL ORNITHOLOGY

PART II

clavicles keep the shoulders apart from each other. The scapular
arch is always present and complete ; the clavicular is sometimes
defective or wanting. There are two leading styles of scapular
arch, corresponding to the ratite and carinate sternum; (1) In
Ratitm the axes of the coracoid and scapula are nearly coincident
(for the most part in a continuous right line) and ankylosed to-
gether ; the clavicles are usually wanting, or defective ; and the

coracoids are instepped on
the sternum far apart. (2)
In all Carinatce the axes
of the coracoid and scapula
form an acute or scarcely
obtuse angle (Figs. 56 and
59, sglc) ; normally these
bones are not ankylosed ;
perfect clavicles are present,
ankylosed with each other,
but free from the other
bones ; and the coracoids
are instepped close together.
Decided exceptions to these
conditions, as in Notornis,
are anomalous ; though in-
completion of the clavicles
repeatedly occurs, as noted
below.

The Copaeoid (Gr. /copaj,
horax^ a crowj ef^os, eidos,
form : the corresponding
bone of the human sub-
ject, which is the stunted
“ coracoid process of the
scapula,” being likened to
a crow’s beak ; no applica-
bility in the present case ;

59 c) is a
stout, straight, cylindric
bone, expanded at each end, extending forward, outward, and
upward from the fore border of the sternum to the shoulder.
Its foot is flattened and splayed to fit in the articular groove
of fore border of the sternum already described ; it often
overlaps that of its fellow on the median line ; is narrower and
remote from its fellow in Ratitce. The head of the bone, irregu-
larly expanded, articulates or ankyloses with the end of the scapula,
and also usually with the clavicle. It bears externally a smooth

Fig. 59. — Right pectoral arch of a bird, Pedicecetes pJia-
sianellus, nat. size, outside view ; Dr. R. W. Shufeldt,

U.S.A. s, scapula; c, coracoid ; p?, glenoid, the cavity
for head of humerus ; d, clavicle ; he, hypoclidium. In
situ, the right end of the figure should tilt up a little ; DiffS 56 C
see Fig. 56. O • J

SEC. IV

ANATOMY OF BIRDS

217

demi-facet, wliich represents the share it takes in forming the
glenoid (Gr. yXrjvTj, glene, a shallow pit ; Fig. 59, gl) cavity, which is
the socket of the humerus. This articular expansion is the glenoid
process of the coracoid : the clavicular process is that by which the
bone unites with the clavicle. The relation between the heads of
the three bones (each uniting with the other two) is such that a
pulley-hole is formed, through which plays the tendon of the pec-
toral muscle which elevates the wing. The coracoid is a very
constant and characteristic bone of birds.

The Scapula (Lat. scapula, the shoulder-blade; Figs. 56, 59, s)
merits in birds its name of “ blade-bone,” being usually a long, thin,
narrow, sabre-like bone, which rests upon the ribs — usually not far
from parallel with the spinal column, and near it ; but in Ratitce
otherwise. It seldom gains much width, and is quite thin and flat
in most of its length ; but it has a thickened head or handle,
expanding outwards into a glenoid process which unites with that of
the coracoid to complete the glenoid cavity, and dilated inward to
form an acromial (Gr. aKpiogtov, akromion, point of the shoulder)
process for articulation with the clavicle (as it does in man), when
that bone exists. The other end is usually sharp-pointed, but may
be obtuse, or even clubbed, as in a woodpecker. The scapula is
broadest and most plate-like in the penguins, in which birds all the
bones of the flipper-like wing are singularly flattened. In Apteryx
it reaches in length over only a couple of ribs ; in most birds,
over most of the thorax ; and in some its point overreaches the
pelvis.

The Clavicles, or Fureulum (Lat. clavicula, a. little key :
furculum, a little fork; Figs. 56, 59, cl), or the clavicular arch, are
the pair of bones which when united together form the object well
known as the “ merrythought ” or “ wish-bone,” corresponding to
the human collar-bones. They lie in front of the breast, across the
middle line of the body, like a V or U ; the upper ends uniting as
a rule both with scapula and coracoid. For this purpose, in most
birds, the ends are expanded more or less ; such expansion is
called the epiclidium (Gr. cTrt, epi, upon ; KXetStov, Ideidion, the
collar-bone) ; in Passerine birds it is said to ossify separately, and
is considered by Parker to represent the procoracoid of reptiles. At
the point of union below, the bones often develop a process (well
shown in the domestic fowl) called the hypoclidium (Gr. vtto, hypo,
under ; Fig. 59, he), supposed to represent the interclavicle of
reptiles. The clavicles are, as a rule, present, perfect, ankylosed
together, articulated at the shoulder ; in a few birds ankylosed
there ; in several, there and with the keel of the sternum ; in
Opisthocomus there and with the manubrium of the sternum. In
various birds, chiefly Picarian and Psittacine, they are defective,

2i8

GENERAL ORNITHOLOGY

PART II

not meeting each other. They are wanting in Struthio, Rhea,
A]pteryx, and some Fsittacidce. Besides curving toward each other,

the clavicles have usually a
fore-and-aft curvature, con-
vex forward. In general,
the strength of the clavicles,
the firmness of their connec-
tions, and the openness of the
V or U, are indications of
the volitorial or natatorial
power of the wings. The
end of the furculum is hol-
lowed for a fold of the wind-
pipe in the crested pintado.

4. THE PELVIC ARCH

The Pelvis (Lat. pelvis, a'
basin, Fig. 60) is that pos-
terior part of the trunk
which receives the urogeni-
tal, and lower portion of the
digestive, viscera. It consists
of the sacrarium on the
middle dorsal line, flanked
on each side by the bones of
the pelvic arch, which sup-
ports the hind limb. In ver-
tebrates generally the pelvic
basin is completed on the
ventral aspect by union {sym-
physis; Gr. (Tvv, sun, together;
cfivo-Ls, growth) of the bones
from opposite sides. Except-
ing only Struthio, which has
a pubic symphysis ; and Rhea,

Fig. 60.— Pelvis of a heron {Ardea herodias), nat. which haS an ischiaC Sym-
size, viewed from below ; from nature by Dr. R. W.

Shufeldt, U.S.A. dl, dorsolumbar vertebrae to and puySlS JUSt DelOW tUe Sacral
including the last one, sc ; below sc, for the extent of vPT*tplvPfP t.hG 'hpIvis of ft

the large black spaces (opposite the arrow) are the VCrteorse, ine pCiVlS OI a
true sacral vertebrae; us, urosacral vertebrae (op- bird IS entirely Open beloW
posite the five oval black spaces ; II, ilium ; Is, i i i i i i •

ischium ; P, pubis ; ob, obturator foramen. The and. behind ; each pelVlC
arrow flies into the acetabulum. ankylosing firmly with

the sacral vertebrae to form a roof over the viscera above
named. This sacro - iliac ankylosis is commonly coextensive
with the confluence of the many vertebrae which make the

SEC. IV

ANATOMY OF BIRDS

219

sacrarium, that is, from the first dorsolumbar to the last
urosacral. The whole roof - like affair looks something like a
keelless sternum inverted. The pelvic arch of each side consists of
three bones, ilium^ ischium, and 'jpuhis, which have independent
ossific centres, but become firmly consolidated together to form the
haunch-bone or os innominatum. Each of these bones unites with
the other two, somewhere near the middle of the whole affair, at a
ring-like structure called the acetabulum (Lat. a vinegar-cruet, Fig.
56, a; Fig. 60, arrow ac), or cotyloid, or coxal cavity, which all
three consequently contribute to the formation of, and which
is the socket for the head of the thigh-bone. When free ribs
issue from under cover of the pelvis, they are commonly ankylosed
with the ilia ; and all the abortive pleurapophyses of the lumbar
and urosacral vertebrae have likewise iliac ankylosis, as explained
in treating of the sacrum. As a whole, the pelvis varies like
the sternum in relative length, breadth, and degree of convexity ;
and especially in the configuration of its posterior border ; but few
zoological characters are de-
rived from this structure.

Viewed from below, the
pelvis is seen to be much
hollowed or excavated for
the lodgment of the kidneys,
and cross-cut into compart-
ments by the sacral rafters ;
the series of sacral bodies
forming a ridge-pole along the middle line. Above, the series of
sacral spinous processes represent the ridge-pole ; anteriorly, the
somewhat spoon-shaped iliac bones are applied, concavity outward,
to the dorsolumbars ; posteriorly, in the middle line, is a more or
less flattened horizontal expansion, and laterally are the more
expanded sides of the ischiac roof, finished along the eaves and
behind by the slender pubic bone, which commonly projects back-
ward, and inclines toward its fellow of the opposite side. The most
prominent formation of the side wall of the pelvis is the thick-lipped
smooth articular ring, the acetabulum, converted in the natural state
into a cup by a membrane. The postero-superior segment of the
rim is prominent, to form the antitrochanter (Gr. dvrt, anti, against ;
Tpoxo^vrrip, trochanter of the femur) against which the shoulder or
trochanter of the femur abuts when the head is in the ring.

It is normal to recent Carinate birds to have the ischium fused
with the ilium, however distinct the pubis may remain ; but to
Cretaceous birds (even Ichthyornis), and the existing Ratitce, to have
both ischium and pubis distinct in most of their extent.

The Ilium (Lat. ilium, haunch-bone ; pi. ilia ; adj. iliac ; Figs.

Fig. 61. — Pelvis of young grouse, showing three
distinct hones. II, Is, P, ilium, ischium, pubis. In
front of former a dorsal vertebra protrudes. (Dr. R.
W. Shufeldt, U.S.A.)

220

GENERAL ORNITHOLOGY

PART II

56, I ; 60, 61, 11) is the median, most anterior and longest of
the haunch-bones, and the only one which extends in advance of
the acetabulum. Such anterior prolongation of this bone is the
specialty of the avian pelvis : it commonly overlies one or more
ribs, and is often overreached by the end of the scapula. It is
longest and narrowest and flattest in some of the lower swimmers;
the reverse among the highest birds. Its relations and connections
have been sufficiently indicated. The bone is almost -always
separated from its fellow by the sacrum, though the approxima-
tion may be close over the back of the pelvis, along the middle line.

The Ischium (Gr. ia-yjiov, ischion, the haunch-bone ; pi. ischia ;
adj. ischiadic, ischiatic, better ischiac ; Figs. 56, 60, 61, Is) lies
entirely post-acetabular, or behind the socket which it contributes
to form, and composes most of the side-wall of the pelvis thence to
the end. It is generally a thin plate-like bone. Among Cretaceous
birds and existing Ratitce it only unites with the ilium at and just
behind the acetabulum, whence a deep ilio-ischiac fissure between
the two exists, as in the grouse. Fig. 61 ; but in ordinary adult
birds this fissure is converted into a fenestra or window of large
size, just behind the acetabulum, by union of the two bones behind
it. This vacuity, whether a notch or a hole, corresponds to the
“ sacro-sciatic notch ” of human anatomy (Fig. 56, in). The ischia of
opposite sides are distinct, except in Rhea.

The Pubis (Lat. pubis, bone of the front of the human pelvis where
the hair grows at puberty ; pi. pubes ; adj. pubic ; Figs. 50, 60, 61, P),
beginning at its share of the acetabular ring, is a long slender bone
which runs along the lower border of the ischium, sometimes for a
short distance only, often for the whole length of the ischium, and usually
projecting behind ; more or less perfectly parallel with, applied to,
or united with, the inferior ischiac border. When separate, a long
deep fissure results ; when united at the end, a long narrow foramen
is formed; when incompletely united in any part of its ischiac
continuity, a fissure and a foramen, in the ostrich two foramina,
result. All these conditions occur ; in any case, such ischio-pubic
interval corresponds to the obturator foramen (Fig. 56,0/ Fig. 60, ob)
of human anatomy ; it is greatest in Cretaceous birds and existing
Ratitce. The free ends of the pubes may be more or less expanded.
In the ostrich only there is a pubic symphysis of the ends of the
bones ; in the same bird a separate ossicle, situated upon the lower
border of the pubes, and called epipubic, is considered to represent
a “ marsupial ” bone (Garrod). In various birds, among them the
ground cuckoo. Geococcyx calif ornianus, the pubis projects a little
forward, under the acetabulum: this prominence is the propubis.
Separation of the pubes is supposed to be for amplification of the
pelvic strait to facilitate the passage of the large eggs birds lay.

SEC. IV

ANATOMY OF BIRDS

221

5. THE SKULL

The Skull of a Bird is a poem in bone — its architecture is the
“ frozen music ” of morphology ; in its mutely eloquent lines may
be traced the rhythm of the myriad amoebiform animals which con-
structed the noble edifice when they sang together.^ The poesy
(TrocrjcrLs, poiesis, a making) of the subject has been translated with
conspicuous zeal and success by Mr. W. K. Parker ; its zoological
moral has been similarly pointed by Professor Huxley; and the
young ornithologist who would not be hopelessly unfashionable
must be able to whistle some bars of the cranial song — the pterygo-
palatine bar at least.

The rapid progress of ossification soon obliterates most of the
original landmarks of the skull, fusing the distinct territories of
bone in one great indistinguishable area. Thus the brain-box of
almost any mature bird is apparently a single solid bone, and most
parts of the jaw-scaffolding similarly run together. Aside from the
bones of the tongue, which are collectively separate from those of
the skull proper ; and of the compound lower jaw, which is freely
articulated with the rest of the skull ; only two or three other
bones of the skull, as a rule, are permanently and perfectly free at
both ends. These are the quadrate bones — the anvil-shaped pieces
by which the lower jaw is slung to the skull ; the pterygoids, arti-
culating the palate with the quadrate ; and sometimes the vomer.
Traces only of distinctness among bones of the face and jaws are
usually found ; but even such vestiges disappear, as a rule, from
among the bones of the brain-box. It is necessary to any intelligent
understanding of the construction of a bird’s skull, to learn some-
what of its mode of development in the embryonic stage ; this being
the only clue to the individual bones of which it is composed, and
so to any correct idea of its morphology. One theory is, that the
skull consists of four modified vertebrae ; and the principal bones
have been named and described by some writers in terms indicating
the elements of a theoretical vertebra. It is true that the skull is
segmented, or may be segmented off, like a chain of several vertebrae;
that it continues the vertebral axis forward; that it has a basis
cranii like a series of vertebral centrums ; above which rises a seg-
mented neural arch enclosing the great nervous mass, and below
which depends a set of bones enclosing visceral parts like a haemal

1 Bone-tissue chiefly consists of the aggregated skeletons of Osteamcebce — a kind
of unicellular protozoan animals which inhabit in myriads the bodies of nearly all
the Vertebrata, possessing the faculty of feeding upon phosphate of lime and other
earthy matters they find in the blood, and afterward excreting them in the form of
multiradiate exoskeletons of their own, collectively forming the whole skeleton of
their host.

222

GENERAL ORNITHOLOGY

PART II

Fig. 62.— Skull of common fowl, enlarged ; from nature by Dr. R. W. Shufeldt, U.S.A. The
names of bones and some other parts are printed, requiring no explanation : but observe the
following points : The distinction of none, of the bones composing the brain-case (the upper
back expanded part) can be found in a mature skull. The brain is contained between the
occipital, sphenoidals, squamosals, parietals, and part of frontal; the ethmoidals belong to the
same group of cranial bones proper. All other bones, excepting the three otic ear-bones, are
bones of the face and jaws. The lower jaw, of five bones, is drawn detached ; it articulates by
the black surface marked articular with the prominence just above— the quadrate hone. Ob-
serve that from this quadrate a series of bones — quadrato-jugal, jugal, maxillary — makes a slender
rod running to the premaxillary ; this is the zygoma, ov jugal har. Observe from the quadrate
also another series, composed oi pterygoid, and palatine bones, to the premaxillary ; this is the
pterygo-palatine har ; it slides along a median fixed axis of the skull, the rostrum, which bears
the loose vomer at its end. The under mandible, quadrate, pterygoid, and vomer are the only
movable bones of this skull. But when the quadrate rocks back and forth, as it does by its
upper joint, its lower end pulls and pushes upon the upper mandible, by means of the jugal
and pterygo-palatine bars, setting the whole scaffolding of the upper jaw in motion. This
motion hinges upon the elasticity of the bones of the forehead, at the thin place just where the
reference-lines from the words “lacrymal” and “ mesethmoid ” cross each other. The dark
oval space behind the quadrate is the external orifice of the ear ; the parts in it to which the
three reference-lines go are diagrammatic, not actual representations ; thus, the quadrate
articulates with a large pro-otic as well as with the squamosal. The great excavation at the
middle of the figure, containing the circlet of unshaded bones, is the left orbital cavity, orhit, or
socket of the eye. The mesethmoid includes part of the background of this cavity, shaded dia-
gonally. The upper one of the two processes of bone extending into it from behind is the post-
frontal or sphenotic process ; the under one (just over the quadrate) is the squamosal process. A
bone not shown, the presphenoid, lies just in front of the oval black space over the end of
hasisphenoid. This black oval is the optic foramen, through which the nerve of sight passes
from the brain-cavity to the eye. The black dot a little behind the optic foramen is the orifice
of exit of a part of the trifacial nerve. The black mark under the letters “ on ” of the word
“ frontal ” is the olfactory foramen, where the nerve of smell emerges from the brain-box to go
to the nose. The nasal cavity is the blank space behind nasal and covered by that bone, and
in the oval blank before it. The parts of the beak covered by horn are only premaxillary,
nasal, and dentary. The condyle articulates with the first cervical vertebra ; just above it, not
shown, is the foramen magnum, or great hole through which the spinal medulla, or main
nervous cord, passes from the skull into the spinal column. The hasioccipital is hidden, ex-
cepting its condyle ; so is much of the hasisphenoid. The prolongation forward of the basi-
sphenoid marked “rostrum,” and bearing the vomer at its end, is the parasphenoid, as far as
its thickened under border is concerned. Between the fore end of the pterygoid and the basi-
sphenoidal rostrum, !is the site of the hasipterygoid process, by which the bones concerned articu-
late by smooth facets ; further forward, the palatines ride freely upon the parasphenoidal
rostrum. In any Passerine bird, the vomer would be thick in front, and forked behind, riding
like the palatine upon the rostrum. The palatine seems to run into the maxillary in this
view ; but it continues on to premaxillary. The maxillo-palatine is an important bone which
cannot be seen in the figure because it extends horizontally into the paper from the maxillary
about where the reference line “maxillary” goes to that bone. The general line from the
condyle to the end of the vomer is the cranial axis, basis cranii, or base of the cranium. This
skull is widest across the post-frontal ; next most so across the bulge of the jugal bar.

SEC. IV

ANATOMY OF BIRDS

223

arch. The hindmost cranial segment, the occipital bone, resembles
a vertebra in many physical characters, and even in mode of
development. But if the serial homology of the skull with the
back-bone be real and true, it is so obscured by the extraordinary
modifications to which the vertebral elements have been subjected
that the fact of such homology cannot be demonstrated ; and to
interpret the skull as something superimposed upon and morpho-
logically different from the spinal column, is perfectly warranted if
not required by the known facts of its constructive development.
This is the view taken by the rulers of to-day’s science. As already

Fig. 63. — Skull of a duck (Clangula islandica), nat. size ; Dr. K. W. Shufeldt, U.S.A. a,
premaxillary bone ; h, partly ossified internasal septum ; V, pervious part of nostril ; c, end of
premaxillary, perforated for numerous branches of second division of the fifth cranial nerve ; d,
dentary bone of under mandible ; e, groove for nerves, etc. ; /, a vacuity between dentary and
other pieces of the mandible ; g, articular surface ; h, recurved “ angle of the jaw ; ” i, occipital
protuberance ; j, vacuity in supraoccipital bone ; k, muscular impression on back of skull ; I is
over the black ear-cavity ; m, postfrontal process ; n, quadrate bone ; 0, pterygoid ; p, pala-
tine ; q, quadratojugal ; r, jugal ; s, maxillary ; t, fronto-parietal dome of the brain-cavity ; u,
the lacrymal bone, immense in a duck, nearly completing rim of the orbit by approaching m ;
V, vomer ; w, supraorbital depression for the nasal gland (see p. 231) ; x, cranio-facial hinge ;
y, optic foramen ; z, etc., interorbital vacuities.

said (p. 202), the relation between cranial and vertebral parts is
rather the analogy of adaptive modification than a true homology
of structure.

Before proceeding to describe the mature skull, it will be best
to consider its mode of development. In this I shall closely follow
Parker, often using the words of that master, and illustrating the
early stages of the embryo with figures borrowed from the same
safe source. In the fewest words possible, I wish to convey an
idea of the embryonic skull up to Parker’s “ third stage,” at which
it begins to ossify. Here, however, I will first insert a figure (Fig.
62), kindly drawn for me by Dr. E. W. Shufeldt, of the U.S. Army,
which shows most of the cranial bones, and will give the student a

224

GENERAL ORNITHOLOGY

PART II

preliminary notion of the “ lay of the land. ” I advise him to con-
template this picture till he has learned the names printed on it by
heart, and can apply them to the identification of the parts of the
real skull he should have in hand at the same time. He may also
meditate on Fig. 63.

Development of the Fowl’s Skull (Figs. 64 to 69). — In the
chick’s head cartilage is formed along the floor of the skull by the
fifth day of incubation. This cartilaginous basilar plate is formed
on each side of the notochord^ Fig 64, c (Gr. vwrov_, noton^ back;

chorde, a chord), a rod-like structure, the primordial axis of
the body, around which, along the spinal column, the bodies of the
vertebrae are formed, and which runs in the middle line of the floor
of the skull as far as the ^pituitary space, pis. The basilar plate is
the parachordal (Gr. irapd, para, by the side of) cartilage. In this,
at the earliest stage, are already planted certain parts of the ear,
the cochlea, cl (Lat. cochlea, a snail-shell), and the horizontal one of
the three semicircular canals, hsc. Opposite the end of the notochord,
the border of the parachordal plate is notched, 5 ; this notch after-
ward forms the foramen ovale, for the passage of parts of the fifth or
trifacial nerve. Near the middle line, posteriorly, the plate is per-
forated for the passage of the twelfth or hypoglossal nerve, g. At
each lateral corner is the separate quadrate cartilage, to form the
quadrate bone. Anteriorly, the plate connects by a strap or bridge
of cartilage, the lingula, Ig (Lat. lingula, a little tongue) with the
trabeculce, tr (Lat. trabecula, a little beam), which enclose the pituitary
space, pts (Lat. pituita, mucus : no applicability here). In front of
this pituitary interval the trabeculae come together to form an inter-
nasal plate, which is so arched over downward as to disappear from
this view, as seen in Fig. 65, where fn is the frontonasal process,
and n is the future external nostril. After uniting in the inter-
nasal plate, the fore ends of the trabeculae separate and become free ;
their free ends are the under extremities of this first visceral arch
(first and only preoral arch).

The same chick’s head, now viewed from below. Fig. 65, shows
the squarish aperture, m, of the future mouth ; the three postoral
arches, with their respective cartilaginous bars, out of which are to
be formed the bones of the jaws and tongue. 1, 2, 3 are the cor-
responding visceral clefts, between the arches ; the first of these is to
be modelled into the ear -passages (outer and middle ear and
Eustachian tube) ; the others will disappear. The quadrate cartilage,
q, is the same that was seen in Fig. 64 ; it is already nearly in
position, between the hind ends of the scaffolding of the upper and
under jaw. The curved subocular or maxillopalatine bar, mxp, de-
veloped in the first postoral arch, already indicates anteriorly
palatine, pa, and posteriorly, pterygoid, pg, parts ; it will form the

SEC. IV

AJVA TOMY OF BIRDS

225

bones so named, and others of the upper jaw. This subocular bar
is an antero-superior part of the first post-oral arch, of which q and
mk are a postero-inferior portion ; the cleft of the future mouth is
to lie between them. The lower jaw-bone, or mandible, is entirely
developed from mk, its several bones developing around this rod of
cartilage, which is called the Meckelian cartilage ; it is to become

Fig. 64. — Skull of chick, fifth day of incu-
bation, X 9 diameters. Seen from above,
the membranous roof of the skull and the
brain removed. cv\, anterior cerebral vesicle ;
e, eye ; c, notochord, running through the
middle of the basilar plate or parachordal
cartilage, in which are already visible the
rudimentary ear-parts, d, the cochlea, hsc,
the horizontal semicircular canal ; pis, the
pituitary space, bounded by tr, the trabe-
culse, which come together before it to form
the frontonasal plate, fn in Pig. 65 ; Ig, lin-
gula or bridge, connecting trabeculse with
parachordal cartilage ; 5, notch afterwards
becoming foramen ovale for passage of parts
of the fifth (trifacial) nerve ; 9, foramen for
hypoglossal nerve ; q, separate cartilage
forming the future quadrate bone. (After
Parker, in Ency. Brit.)

Fig. 65. — Same as Fig. 64, but seen from below,
cwl, anterior cerebral vesicle ; e, eye ; m, mouth ;
pis, pituitary space ; fn, frontonasal plate ; tr,
ends of the trabeculae, free again after their union
and bent strongly from the original axis of the
trabeculae ; n, external nostril ; mxp, subocular
bar of cartilage, or pterygopalatine rod, to form
pa, palatine, and pg, pterygoid bone, and other
parts of the upper jaw, as the maxillary, jugal,
and quadra tojugal ; q, quadrate cartilage, same
as seen in Fig. 64 ; mk, Meckelian cartilage, to
form lower jaw — these parts are in the first
postoral visceral arch ; cJi, ceratohyal, and hh,
basihyal, of second postoral arch ; cbr, cerato-
branchial, ebr, epibranchial, bbr, basibranchial,
of third postoral arch ; the parts of the second
and third arch all going into the hyoid bone. 1,
2, 3, 1st, 2d, 3d visceral clefts, whereof the 1st is
to be modified into the ear-passages and the
others are to be obliterated. (After Parker.)

movably articulated with the bone, the quadrate, into which q will
be transformed. Thus the postero-inferior part of the first postoral
arch (second of the whole series of arches) begins in two pieces, one
of which is to become the suspensorium, or suspender of the mandible,
and the other the mandible itself. The rest of the pieces belong to
the second and third postoral arches, and all together make up the
very composite hyoid bone, or bone of the tongue (Figs. 72, 73, 74).
The pieces ch and hh are in the second arch, and form respectively
Q

226

GENERAL ORNITHOLOGY

PART II

the ceratoliyal and basihyal bones ; the pieces chr, ebr, and bbr are in
the third arch, and form respectively the ceratobmnchial, epibranchial,
and basibranchial bones. These pieces of the third arch have already
outgrown those of the second arch, and they will form the greatest
part of the hyoid bone.

In the second stage, after the fifth day of incubation, but before
any ossification has begun, a vertical section shows the appearances
represented in Fig. 66. The parachordal and trabecular cartilages

are applied to each

other unconformably,
the latter rising high
between second and
third cerebral vesicles
to form the posterior
pituitary wall, pci, in
which the axial skele-
ton properly ends.
There are other

changes in the para-

chordal cartilages.
The internasal plate,
formed by the union
of the trabeculae in
front of the pituitary
space, has become a

Fig. 66. — Head of a chick, second stage, after five days of l ii

incubation, section in profile ; X 6 diameters. ctI, ci;2, cv3, first, Vertical meclian Wall

second, and third cerebral vesicles ; 1, place of the first nerve, 'kpfwfia'n ftia rklfc»pfrkT-i7-

the olfactory ; 2, place of second nerve, the optic ; ic, internal 6iie UiicU.bUiy

carotid artery, running into skull at what was originally the and Optic chambers
pituitary space, now an opening bounded in front by the an- r i r i j. j i ri.

terior, ad, behind by the posterior, pd, clinoid walls ; nc, noto- 01 the right and left
chord; oc, occipital condyle, thence to pd being the original

parachordal cartilage, here seen in profile ; eo, exoccipital ; eth, ®m.es \pn ailU tiiL, to

■ethmoid, with ps, its presphenoid region posteriorly, and pn, rpg and alc^. This

prenasal part ; this whole plate afterward developing into parts . . '

of the nose and the partition between the eyes ; pa, palatine ; partition, bosidos
pg, pterygoid region ; pa and pg reference lines are in the o • -R 11 +V,

chick’s mouth ; mfc, Meckelian cartilage (lower jaw) : cTi- and M, lOrmillg Unally the

(TfSfplrker”? intcTorbital septum

which divides the

right and left orbits, will undergo further notable changes
in direction, and will develop lateral plates and processes, which
will make up the nasal labyrinth and the partition between the
■cavity of the nose and that of the eye, when any exists. Such
lateral developments of the ethmoid plate are the aliethmoid, aliseptal,
and alinasal. This plate extends backward in mid-line to the optic
foramen, 2, ending in the anterior clinoid wall, acl, separated from the
(parachordal) posterior clinoid, pci, wall by the original pituitary
space, now the opening through which the carotid arteries, ic, enter
the brain cavity. Besides ethmoidal parts proper, the plate

SEC. IV

ANATOMY OF BIRDS

227

develops at what will be the end of the upper beak a p'enasal car-
tilage, 'pn, to become the axis of the beak. The mouth is become
already better formed, the axis of its cavity pointing more forward
than downward ; and great changes are undergoing in parts of the
ear at the back corner of the mouth. The quadrate and Meckelian
cartilages are assuming much of their true form. The quadrate
develops an orlital process which extends free into the orbit, and an
otic process which articulates with the auditory sac and parts of the

exoccipital cartilage. The rela-
tions at this stage have not been
made out in the fowl, but are
figured and described from the
corresponding stage of the Eu-
ropean house -martin {Chelidon
urUca). In Fig. 67, mk is the
cut stump of the Meckelian carti-
lage, of which ar is the articular
part ; g is the quadrate, of which
a backward process is seen arti-
culating with teo, the tympanic
wing of the exoccipital. Just
below and behind this otic pro-
cess of the quadrate, exactly
where in riper embryos is the
fenestra ovalis in which is fitted
the foot of the stapes or stirrup-
bone of the middle ear, there
appears a trowel-shaped projec-
tion of cartilage, the handle of
which is continuous with the
substance of the ear-capsule ; the
sickle-shaped piece behind which
is the tympanic wing of the ex-
occipital (teo). This trowel of
cartilage is the upper anterior seg-
ment of the hyoidean (second post-

Fig. 67. — The postoral arches of the house
martin, at middle of period of incubation, la-
teral view, X 14 diameters, mlc, stump of Mec-
kelian or mandibular rod, its articular part, ar,
already shapen ; q, quadrate bone, or suspen-
sorium of lower jaw, with a free anterior orbital
process and long posterior otic process articu-
lating with the ear-capsule, of which teo, tym-
panic wing of occipital, is a part ; mst, est, sst,
ist, sth, parts of the suspensorium of the third
postoral arch, not completed to chy; mst,
mediostapedial, to come away from teo, bring-
ing a piece with it, the true stapes or columella
auris ; the oval base of the stapes fitting into
the future fenestra ovalis, or oval window look-
ing into the cochlea ; sst, suprastapedial ; est,
extrastapedial ; ist, infrastapedial, which will
unite with sth, the stylohyal ; chy and hhy,
ceratohyal and basihyal, distal parts of the
same arch ; bbr, hr 1, hr 2, basibranchial, epi-
branchial, and ceratobranchial pieces of the
third arch, composing the rest of the hyoid
bone ; tg, tongue. (After Parker.)

oral) arch, being to that arch what the pterygo-palatine bar is to the
mandibular (first postoral) arch. Several parts of this stapedial
cartilage are recognised, as named in the fine print under the figure.
If the connections of the second postoral arch were completed, as
those of the first are, the tongue bone would be slung to the skull
as the lower jaw is ; but they are not, the tract represented by the
dot-line from the stylohyal, sth, to the ceratohyal, chy, being, like ist,
above sth, only soft connective tissue. This defect of connection is
made up for by the great development of the hyoidean parts of the

228

GENERAL ORNITHOLOGY

PART II

third post-oral arch, If 1 and hr 2, which retain the tongue-bone in
position, without however articulating it with the skull. The hand
of the trowel of cartilage soon segments itself off from the ear-
capsule, bringing away with it a small oval piece of the periotic wall,
which piece is the true stapes, and the oval space in which it fits is
the fenestra ovalis leading into the inmost ear (the cochlea). The
broad part of the trowel-blade is the extrastapedial part, on which
the memhrana tympani, or ear-drum, will be stretched. The stylo-
hyal, sth, will join the extrastapedial plate, and the afterward

chondrified band of union will
be the infrastapedial, ist. (Figs.
71, st, and 83.)

Returning now to the chick’s
head, which we left to examine
the intricate ear-parts at the
proximal end of the second
post-oral arch, we see by Fig.
68 how rapidly the parts are
shaping themselves at the end
of this second stage of develop-
ment. This figure shows the
cartilaginous skull, in which
no trace of ossification has
appeared, excepting in the
under mandible. The brain
and membranous parts of the
cranium have been removed.
The roof of the skull never
becomes cartilaginous, bone
there growing directly from
the membrane ; and the whole
of the chondro - cranium, as
shown in the figure, is one continuous cartilaginous structure (like the
whole skull of an adult shark or skate), excepting the parts of the post-
oral arches, which are separate. The auditory capsule is environed
by occipital cartilage, eo, stretching over the back of the skull, and
by wing-like growths {alisphenoids, as) which wall most of the brain-
box in front. The high orbito-nasal septum is a continuous vertical
plate of cartilage, upgrowing from the tract of the conjoined tra-
beculae. Lateral developments of this ethmoidal wall, in front, are
divided into several recognisable parts, ale, als, aln, the latter being
the external nostril ; is a transverse partition between the orbital
and nasal chambers. The nasal cartilages ultimately become much
convoluted to form the nasal labyrinth, among the convolutions of
which will be the superior and inferior turbinal cartilages, in addi-

Fig. 68. — Skull of chick, second stage, in profile,
brain and membranes removed to show cartilagi-
nous formations, x 4 diameters, eth, ethmoid,
forming median nose-parts and interorbital sep-
tum ; developing lateral parts, as ale, aliethmoid,
als, aliseptum, aln, alinasal, pp, partition between
nose and eye ; pn, pre-nasal cartilage ; ps, pre-
sphenoidal part of mid-ethmoid ; 2, optic foramen ;
as, alisphenoid, walling brain-box in front ; pj,
post-frontal, bounding orbit behind ; pa, pg, pala-
tine and pterygoid ; q, quadrate ; so, supra-
occipital ; eo, exoccipital ; oc, occipital condyle,
borne upon basioccipital, and showing no, remains
of notochord ; these occipitals bound the foramen
magnum, and eo expands laterally to form a tym-
panic wing, circumscribing the external auditory
orifice behind and below ; Use, psc, horizontal and
posterior vertical semicircular canals of ear ; fr,
st, fenestra rotunda and fenestra ovalis, leading
into inner ear, latter closed by foot of the stapes ;
Ttik, ch, bJi, hbr, cbr, ebr, parts of jaw and tongue,
as named in Pigs. 65, 66, and 67. (After Parker.)

SEC. IV

ANATOMY OF BIRDS

229

tion to those already noted. The ethmoidal wall ends behind at
the presphenoidal region, where the brain-case begins ; below
and behind, it is deeply notched for the o-ptic foramen, 2. The
pituitary space forms a circular foramen, through which the carotid
arteries enter. The site of the orbit of the eye is bounded behind
and below by the post-frontal process of the alisphenoid wing, pf of
as. The pterygopalatine rod is seen along the under border of the
skull, pg and pa. The quadrate, q, has acquired nearly its shape,
and the rest of the mandibular and hyoidean parts are clearly dis-

Fig. 69.— Skull of chick, third stage,
viewed from helow, x 6f diameters, pn,
prenasal cartilage, running behind into the
septum nasi ; on each side of it the premax-
illary, px, of which the (inner) palatal and
(outer) dentary processes are seen (the upper
nasal process hidden) ; mx, the maxillary,
developing inner jjrocess, the maxillopala-
tine, mxp ; pa, the palatal, well formed,
articulating behind with rbs, the sphenoidal
rostrum, its thickened under border, the
parasphenoid ; this will bear the vomer at
its end Avhen that bone is developed ; j,
jugal, joining mx and qj, the quadratojugal,
joining j and q, the quadrate ; mx to q, the
jugal bar or zygoma; pg, the pterygoid,
making with pa the pterygopalatine bar,
joining q and px ; ht, the basitemporal, great
mat of bone from ear to ear, underflooring
the skull proper, as rbs, a similar formation,
does further forward ; ic, outer end of
carotid canal, to run between the bt plate
and true floor of skull, and enter brain
cavity at original site of pituitary fossa
(Figs. 64, 66, ic) ; ty, tympanic cavity— ex-
ternal opening of ear ; as, alisphenoid,
bounding much of brain-box anteriorly, and
orbital cavity posteriorly ; psc, posterior
semicircular canal of ear, in opisthotic bone,
which will unite with the spreading eo, ex-
occipital, which will reach the condyle
shown in the middle line, above the foramen
magnum, /m, completed above by so, supra-
occipital ; 8, foramen lacerum posterius,
exit of pneumogastric, glossopharyngeal
and spinal accessory nerve ; 9, exit of hypo-
glossal nerve, in basioccipital. (After
Parker.)

played, mk, etc. The proximal hyoidean element, st, is freed from
the periotic cartilage, leaving the fenestra ovalis (see last paragraph).
Below the general outline, pa to oc, is not shown a mat of soft
tissue, in which are to be developed the basitemporal and parasplien-
oid bones which underfloor the whole skull, — the former making a
plat between the ears, Fig. 69, ht, the latter forming the thickened
under edge of the rostrum of the skull rbs.

At the third stage, about the middle of the second week of in-
cubation, the cartilaginous parts already described are neatly finished,
and the skull is beginning to ossifij. The occipital parts are well

230

GENERAL ORNITHOLOGY

PART II

formed ; the condyle is perfect ; the foramen magnum is circum-
scribed by the ex- and supra-occipitals, eo and so, Fig. 69. Investing
bones, formed in membrane without previous cartilage, are becom-
ing apparent. The basitemporal, ht, and parasphenoid, rhs, are en-
grafting upon the base of the skull. The prenasal cartilage, pn, now
at its full growth, is beginning to decline ; on each side of it is
formed a three-forked bone, the premaxillary, px, having superiorly
nasal, and laterally palatal and dentary processes. This bone is to
grow to great size, forming most of the upper beak, and starving
out the maxillary, which in mammals is the principal bone of the
upper jaw. The palatal, pa, and pterygoid, pg, bones are ossified,
and the quadrate, g, is ossifying. Between the premaxillary and
the quadrate are the bones forming the zygoma, or jugal bar, de-
veloped in the outer part of the maxillopalatine bar of the earlier
embryo. They are the weak maxillary, mx, with its ingrowing
process, the maxillopalatine bone, mxp ; next the jugal, j ; then the
quadratojugal, qi ; the whole forming an outer lateral rod from
quadrate to premaxillary, like a duplicate of the pterygopalatine
rod from the same to the same.

Among occurrences of later stages are to be noted the development
in membrane in the middle line below of the vomer, borne upon the
end of the rostrum ; the roofing in of the whole skull by the parietal,
squamosal, frontal, and nasal bones ; the completion of the periotic
hones as the prootic, epiotic, and opisthotic, which form the otic capsule,
or otocrane ; the development of lacrymal hones, bounding the orbits of
the eyes in front. Absorption of the middle wall of cartilage be-
tween the nasal and orbital cavities nicks off the nose parts from
those of the orbit (Fig. 70, between nth and eth) ; and certain changes
in the orbital septum develop the orhitosphenoids. Very nearly all
the bones of a bird’s skull having thus been accounted for, we may
next consider them in their adult condition. Eeference should
be made to Figs. 62, 63, 70, 71.

The Occipital Bone (Figs. 62, 70, 71) forms the back part of the
floor of the skull, and lower part of the back wall of the skull ;
neither its boundaries nor its composition are visible in adult skulls.
It is formed by the hasioccipital, ho, below in the middle line ; the
supraoccipital, so, above in the middle line ; the exoccipital, eo, on either
side. These bound th.Q foramen magnum (Fig. 69, fm), where the
nerve-mass makes its exit from the cavity of the cranium into the
tube of the spinal column. At the lower part of the foramen is the
protuberant occipital condyle (Figs. 68, 71, oc), borne chiefly upon
the hasioccipital, but to the formation of which the exoccipitals also
contribute ; the latter flare widely on each side, into the tympanic
wings, which bound the external auditory meatus behind. The true
hasioccipital is mostly covered by the underlying secondary bone.

SEC. IV

ANA TOMV OF BIRDS

231

the hasitemporal (Figs. 69, 70, U), which extends from one tympanic
cavity to the other, and more or less forward in the middle line to
the sphenoidal rostrum. Openings to be observed in the occipital
region, besides the great foramen, are those for the hypoglossal
nerve, 9, near the condyle ; for the parts of the vagus nerve, 8,
more laterally, and the carotid canal, ic: also, above the foramen
magnum, openings for veins, sometimes of great size, as in Fig. 63,/.

The Parietals (Figs. 62 and 70, 71). — Proceeding up over

the brain-box, the next bones are a pair of parietals between the
occipital behind, the frontal before, and the squamosal beside ; but
their limits are rarely if ever to be seen in adult skulls. They are
relatively small in birds ; simply squarish plates, bounded as said,
coming together in the upper mid-line, or sagittal suture.

The Frontals (Figs. 62 and 70,/, 71), originally paired, soon fuse
together, and with surrounding bones of the skull, though main-
taining some distinction from those of the nose and jaw. These
roof over much of the brain cavity, close in much of it in front, and
form the roof and eaves of the great orbital sockets. Anteriorly
in the middle of the forehead line the feet of the nasal process of
premaxillary are implanted upon the frontal, usually distinctly ;
more laterally, the nasal bones are articulated or ankylosed ; this
fronto-naso-premaxillary suture forming the frontofacial hinge (Fig.
63, x), by the elasticity or articulation of which the upper jaw
moves upon the' skull, when acted on by the palatal and jugal bars.
In the midst of the forehead the two halves of the frontal some-
times separate, as they do in the fowl, allowing a little of the
mesethmoid to come to the front. In the middle line, underneath,
the frontals fuse with whatever extent there may be of the meseth-
moid which forms the lengthwise interorbital septum, and often a
crosswise partition between the orbital and nasal cavities. To the
antero-external corners of the frontal are articulated or ankylosed
the lacrymals. The postfrontal process} morphologically the post-
frontal or sphenotic bone, bounds the rim of the orbit behind ; it
is usually quite prominent. The frontal rim of the orbit in many
birds shows a crescentic depression (very strong in a loon and many
other water-birds; Fig. 63, w), for lodgment of the supraorbital
gland, the secretion of which lubricates the nasal passages. The
cerebral plate of the frontal is often imperfectly ossified, showing
large fenestrse besides the regular openings for the exit of nerves

^ There is apparently some ambiguity in the use of the term “postfrontal”
process by different authors. It would appear that this process, bounding the rim
of the orbit behind, may be a projection of the frontal bone, and therefore strictly a
postfrontal process. Or that, as said by Owen for Rhea, it may be a separate bone,
and therefore properly a postfrontal hone. Or, again, that it may have nothing to
do with the frontal bone, but belong to the alisphenoid, as a process of the latter or
a separate ossification ; in which case it would be properly the sphenotic. In no
event has it anything to do with the squamosal process lettered as such in Fig. 62.

232

GENERAL ORNITHOLOGY

PART II

which are always found at the back of the orbit. Viewed from above,
the frontal is vaulted and expanded behind, over the brain cavity,
then pinched more or less, sometimes extremely narrow over the
orbits, then usually somewhat expanded again at the frontofacial
suture. The extent of the frontal between the orbits and face, in
the lacrymal region, is very great in the duck family, as see in Fig. 63.

The Squamosal (Lat. squama^ a scale ; Figs. 70, 71, sq) bounds
the brain -box laterally, between occipital, parietal, frontal, and
sphenoidal bones, its distinction from all of these being obliterated
in adult life. It is situated near the lower back lateral corner of
the skull, forming some part of the cranial wall just over the ear-
opening, and a strong eaves for that orifice. It is firmly united also
to the bones of the ear proper, and receives the larger share of the
free articulation which the quadrate has with the skull. It often
develops a strong forward-downward spur, the squamosal process
(Fig. 62), looking like a duplicate postfrontal process; between
these two is the crota^phite depression, corresponding to the “tem-
poral fossa ” of man, in which lie the muscles which close the jaws.
It scarcely or not enters into the orbit, the adjacent part of the
orbit being alisphenoidal.

The Periotic Bones (Gr. Trepl, peri, about ; ovs, wros, ous, otos,
the ear ; Fig. 70) are those that form the petrosal hone (Liit. petrosus,
rocky, from their hardness), or bony periotic capsule, containing the
essential organ of hearing. When united with each other and with
the squamosal, they form the very composite and illogical bone
called “ temporal ” in human anatomy. There are three of these
otic bones, — an anterior, the prootic ; a posterior and inferior, the
opisthotic (Gr. oTna-Oe, opisthe, behind) and a superior and external,
the epiotic. They can only be studied in young skulls, upon careful
dissection ; they do not appear upon the outside of the skull at all,
excepting a small piece of the opisthotic, which there fuses indis-
tinguishably with the exoccipital. But some part of these bones
is seen on looking into the cavity of the outer ear, and if the
fenestra ovalis can be recognised, it determines a part of the bound-
ary between the prootic and opisthotic bones, while the fenestra
rotunda lies wholly in the latter. The cavity of the periotic bone
is hollowed for the labyrinth of the internal ear, including the
cochlea, which contains the essential nervous organ of hearing, and
the three semicircular canals — so much of them as does not invade
surrounding bones. In the young fowl’s skull viewed internally (Fig.
7 0), Parker figures a very large prootic portion (po) of the periotic,
perforated by the internal auditory meatus (7) for the entrance from
the brain of the auditory nerve ; below and behind the prootic a
small opisthotic (op), in relation with the exoccipital, upon the
surface of which it also appears, outside (Fig. 69, at ^sc), and with

SEC. IV

ANATOMY OF BIRDS

233

which it blends ; a very small epiotic centre (e^), between the
prootic and supra-occipital ; and the anterior semicircular canal (asc)
embedded in the latter. In Dr. Shufeldt’s figure the otic elements
are merely noted diagrammatically. According to Huxley’s gener-
alisation, the epiotic is in special relation with the posterior semi-

Fig. 70. — Ripe chick’s
skull, longitudinal section,
viewed inside, x 3 diameters;
after Parker. In the man-
dible are seen ; mk, remains
of Meckelian rod ; d, dentary
bone ; sp, splenial ; a, an-
gular ; su, surangular ; ar,
articular ; iap, internal arti-
cular process ; pap, pos-
terior articular process. In
the skull : pn, the original
prenasal cartilage, upon
which is moulded the pre-
maxillary, px, with its nasal
process, npx, and dentary
process, dpx ; sn, septo-
nasal cartilage, in which is
seen nn, nasal nerve ; nth,
nasal turbinal ; the refer-
ence line crosses the cranio-
facial suture, the face parts
and cranial parts being
nearly separated here by
the nick seen in the ori-
ginal cartilaginous plate ;
eth, ethmoid ; pe, perpen-
dicular plate of ethmoid,
which will spread nearly
throughout the dotted car-
tilaginous tract in which it
lies, to form nearly all the
interorbital septum ; trans-
verse thickening (in some
birds) below the reference
line eth will form the pre-
frontal or orbitonasal sep-
tum ; iof, interorbital fora-
men ; ps, presphenoidal
region, just above which is
the orbitosphenoidal region ;

2, optic foramen ; as, ali-
sphenoid, with 5, foramen
for divisions of the 5th
(trifacial) nerve ; /, frontal ;
sq, squamosal ; p, parietal ;
so, superoccipital ; asc, an-
terior semicircular canal ;
sc, a sinus (venous canal) ;
ep, epiotic; eo, exoccipital I op, opisthotic; po, prootic, with 7, meatus auditorius internus,
for entrance of 7th nerve ; 8, foramen for vagus nerve ; ho, basioccipital ; ht, basitemporal ; ic,
canal (in original pituitary space. Fig. 66, ic) by which carotid artery enters brain cavity ; ap,
basipterygoid process ; ap to rhs, rostrum of the skull, being the parasphenoid bone under-
flooring the basisphenoid and future perpendicular plate of ethmoid. (The scaffolding of the
upper jaw not shown, excepting px, etc.)

circular canal ; the prootic with the anterior vertical canal, between
which and the foramen ovale (5) for the lower divisions of the
trifacial nerve it lies. That part on which the inner foot of the
quadrate is implanted is prootic. Below the drooping eaves of the
squamosal, before the flaring wing of the exoccipital, and behind
the quadrate bone, is the always decided and considerable cavity of

234

GENERAL ORNITHOLOGY

PART II

the ear, bounded pretty sharply by the squamosal and exoccipital
rim, sloping with less distinction in front toward the orbital cavity.
In this auditory hollow may be seen several openings : the meatus

or proper ear-
passage, through
which, in one direc-
tion, a bristle may
be passed to emerge
at or near the
middle line of the
base of the skull,
about the root of
the basisphenoidal
rostrum. Such a
passage is through
the first msceral
cleft of the early
embryo, modified
into meatus audi-
torius and Eusta-
chian tube, which
latter communicates
with the back part
of the mouth. Be-
sides the other ear-
passages proper,
may be found other
openings of air-
passages leading
into the interior
diploic tissue of
bones of the skull.

Fig. 71.— Ripe chick’s skull, in profile, x 3 diameters ; after and especially into
Parker, px, premaxillary; aln, alinasal cartilage; en, septo- T

nasal ; n, nasal bone ; I, lacrymal ; pe, perpendicular plate of ^ne lOWCr jaW-DOnC.
ethmoid, as in Fig. 70 ; ps, presphenoidal region ; as, alisphe- T’Rp
noid ; /, frontal ; p, parietal ; sq, squamosal ; so, superoccipital ;
eo, exoccipital ; oc, occipital condyle ; st, the cross-like object, immensely
the stapes, whose foot fits fenestra ovalis, see Fig. 83 ; q, quad- •, J ^

rate ; pg, pterygoid ; qj, quadratojugal ; j, jugal ; pa, palatine ; lOpeU in OWlS, in
mx, maxillary. In the mandible: d, dentary ; su, surangular ; Q-nppiPS of

a, angular ; ar, articular ; iap, internal angular process ; pap, bpeoitJb ui

posterior angular process. 2, optic foramen ; 5, foramen ovale, "which thev Ure Un-
for inferior divisions of the 5th nerve. (Compare Fig. 70.) . ^

symmetrical, that

is, not sized and shaped alike on right and left sides of the head.

The Sphenoid (Gr. o-chyv, sphen, a wedge ; etSo?, eidos, form ;
Figs. 62, 70, 71) is a compound bone, not easy to understand as it
occurs in birds, as much of it is hidden from the outside, some of
it is very slightly developed, and all of it is completely consolidated

ear-parts are
deve-

SEC. IV

AJVA TO MY OF BIRDS

235

with surrounding bones in the adult. It is wedged into the very
midst of the cranial bones proper, with its body in the middle line
below, next in front of the basioccipital, and its wings spread on
either side in the orbital cavity. A sphenoid consists essentially
of the lasisplienoid, or main part of the bone (Fig. 62) ; the pair
of alisphenoids or “wings,” one on either side (Figs. 70, 71, as);
the obscure presphenoid {ps) in the middle line in front of and
above the main body ; and the pair of small orUtosphenoids,
which are in fact the wings of the presphenoid. The body is
usually covered in by the underflooring of the basitemporal ; it
is a flat triangular plate, produced more or less forward in the
middle line as the basisphenoidal rostrum, or beak of the skull.
This rostrum {ap to rbs in Fig. 70) is an important thing. It forms,
in fact, the central axis of the base of the skull ; with the meseth-
moid plate the inferior border of the interorbital septum, usually
thickened by the underflooring of the parasphenoid (Fig. 70, rhs).
The rostrum often bears on each side a hasipterygoid process {ap), — a
smooth facet with which the pterygoid articulates. These processes
may be very strong, and far back on the basisphenoid body, when
the pterygoids articulate with them near their own posterior ends,
as in the struthious birds and tinamous (Fig. 75, Up); or they may
be further along on the rostrum, and the pterygoids then articulate
near or at their fore -ends. The rostrum may be produced far
forward, beyond the maxillopalatines and vomer even, as in an
ostrich ; or it may bear the vomer at its end ; or may be embraced
by forks of the vomer ; the palatines may glide along it, or be
remote from it on either side. In any event, whatever its produc-
tion, whatever part may be ethmoidal, or basisphenoidal, or para-
sphenoidal thickening, ptery go-faceting, etc., this “beak” of the
basisphenoid is always in the axis of the base of the skull, and
at the bottom of the interorbital plate ; it may be horizontal, or
obliquely ascending forward ; and the variety of its relations with
the pterygopalatine and vomerine mechanism furnishes important
zoological characters, as we shall see when we come to treat of
palatal structure particularly. Just at the base of the beak, where
it widens into the main body of the bone, may commonly be seen,
coming from between the sphenoidal body and the lip of the basi-
temporal underflooring, the orifices of the Eustachian tubes, and
often also the anterior ends of the carotid canal. If a bristle,
passed into a questionable foramen here, comes out of the ear, it
has gone through the Eustachian tube ; if it comes out below the
ear, on the floor of the skull, outside, it has run in the carotid
canal. The extent of the alisyhenoids (Figs. 70, 71, as) cannot
be determined in old skulls. They lie at the back lower border
of the orbital cavity, closing in most of the brain-box that is

236

GENERAL ORNITHOLOGY

PART II

not foreclosed by the frontal bone. You will always find at the
back of the orbit, close to the mid-line, and rather low down,
the very large holes which transmit the optic nerves from the
brain to the eye ; these are the o'ptic foramina (any Figs., 2) ;
alisphenoid should not extend in front of these orifices. A
little below and behind the optic foramina, and much more
laterally, not far from the quadrate itself, is a considerable fora-
men, quite constant, for transmission of the inferior divisions of
the fifth {trigeminal or trifacial) nerve. This is the foramen ovale
(any Figs., 5) ; it is either in the alisphenoid, or between that bone
and the prootic j it must not be mistaken for one of the several
smaller holes, usually seen close about the optic foramen, which
transmit the three nerves {oculimotor, ‘pathetic, and abducent) which
move the muscles of the eyeball; these holes being collectively
about equivalent to the foramen lacerum anterius of human anatomy.
Parts about the optic foramen, before and above, are presphenoidal
(Figs. 70, 71, ps) and orbitosphenoidal ; but they are obscure to
all but the embryologist, and furnish no zoological characters.

The Ethmoid (Gr. -gOpos, ethnos, a sieve ; from the way it is
perforated in the human species ; Fig. 62) is the bone of the mid-
line of the skull, in front of the sphenoidal elements and below the
frontal ; it is in special relation with the olfactory nervous apparatus,
or sense of smell. This is not an easy bone to “ get the hang of ”
in birds. Eeferring to Figs. 66, 68, eth, the student will see in the
early embryo a high thin plate of cartilage, the mesethmoid car-
tilage, which is developing lateral processes to form the convoluted
walls of the nasal passages. By the uprising and forth-growing of
the prenasal cartilage, the mesethmoidal plate is tilted backward,
as it were, under the frontal. Next, by absorption of tissue just
opposite the future craniofacial suture, the plate is nicked apart,
the portion in front of the nick elaborating the nasal chambers,
which usually remain cartilaginous, and the portion behind this
nick becoming the permanent plate. Fig. 7 0, eth, pe, to which the
name mesethmoid or mid-ethmoid is more strictly applicable. Prac-
tically, a bird’s ethmoid is chiefly the interorbital septum, in vertical
mid-line between the orbits, with such flange-like processes or lateral
plates as may be developed to form an orbitonasal septum separating
the eye-socket from the nose-chamber. In general, the permanent
ethmoidal plate becomes nearly coincident with this orbital wall,
and pretty well cut off from the osseous or cartilaginous develop-
ments, when any, in the nasal cavities. It is then fairly under
cover of the frontal, with which, as with the sphenoidal elements
posteriorly, it becomes completely fused. When this interorbital
septum is fully developed, it completely divides the right and left
orbital cavities, and its lower horizontal border, fused with the

SEC. IV

ANA TOMY OF BIRDS

237

basisphenoidal rostrum, may like the latter be thickened by bearing
its share of the parasphenoidal splint. Oftener, however, this lower
border slopes upward and forward, from the sphenoidal base to the
roof of the skull about the site of the craniofacial suture ; and
usually the septum is incomplete, having a membranous fenestra
somewhere near its middle (Fig. 70, iof). Along the upper border
of the mesethmoid plate, or just in the crease between it and the
overarching frontal, may usually be seen a long groove, which, be-
ginning behind at the olfadonj foramen of the brain-box, conducts
the thence -issuing olfactory nerve to the nasal chambers. Some-
times there is another such groove, from a foramen near by in
the sphenoidal parts, which similarly traces the course of the
ophthalmic (first) division of the trifacial nerve. Occasionally, as
in the fowls, the two halves of the frontal bone separate a little at
the extreme forehead, allowing the mesethmoid plate there to come
up flush with the outer surface of the skull.

In some birds, as the low ostrich, for example, the original
mesethmoidal cartilage-plate does not nick apart into orbital and
nasal moieties, but ossifies as a continuous sheet of bone, dividing
right and left halves of the skull far towards the point of the beak
(see Fig. 75, beyond Pv to Pmx). A nasal septum, separated from the
orbital septum, may persist to ossify ; forming, as in the raven, a
vertical plate separate from all surroundings, and liable to be mis-
taken for a free wmer (see Fig. 79, where the reference line v goes
to it, instead of to the truncate vomer) j or, as in many birds, a
plate variously ankylosed with its surroundings. But these forma-
tions, as well as the various turlinal (Lat. turbo, a whorl) scrolls and
whorls formed in this part of the skull, belong rather to the organ
of smell than to the skull proper.

The Cranial Bones proper are all those thus far described,
excepting the nasal ossifications just noted, which belong to the first
preoral arch ; and the stapedial parts of the ear, which belong to
the hyoidean apparatus (second postoral arch). Intermediate in
some respects between the proper cranial bones and

The Facial Bones proper is the Vomer. — By “ facial bones,”
as distinguished from “cranial” bones, is meant the entire bony
scaffolding of the upper and lower jaws, and of the tongue, —
parts developed in the preoral or maxillary, and first, second, and
third postoral, or mandibular, hyoidean proper, and branchial,
arches.

The Vomer (Lat. wmer, a ploughshare ; Figs. 62, 63, 75 to 80, «^)
vras considered, by those who held the vertebral theory of the skull,
to be the body of the foremost (fourth from behind — the basioc-
cipital, basisphenoid, and presphenoid being the other three) cranial
vertebra. So far from having any such morphological significance.

238

GENERAL ORNITHOLOGY

PART II

it is one of the late secondary bones, developed, if at all, apart from
the general make-up of the skull, as a special superaddition under-
lying the ethmoidal region, as the parasphenoid and basitemporal
underlie the skull farther back. Its character is extremely variable
in the class of birds, though usually constant in the several natural
divisions of the class, — a fact which confers high zoological value
upon this anomalous bone. A vomer is a symmetrical mid-line
bone of the base of the skull, found if at all at or near the end of
the rostrum. It is originally double, i.e. of right and left paired
halves. These halves persist distinct in the woodpeckers, and are
remote from each other, one on each side of the mid-line (Fig. 80).
The vomer is wanting entirely in the Columbine birds, as the
pigeons and some of their allies, as the sand-grouse {Pteroclidce) and
bush-quails {Uemipodiidce) of the old world, and in certain of the
true Gallinm. Its connections are various. It may be borne free
upon the end of the rostrum. It may be applied like a splint by
a grooved upper surface to the under side of the rostrum, and so
fixed there ; or, in such situation, it may glide along the rostrum
according to the movements of the palatal j^arts with which it may
connect. Thus, in the ostrich (Fig. 75), it saddles the rostrum
below, and is joined by the maxillopalatines. Or, it may be united
with separate ossifications, the septomaxillaries, which in some birds
bridge across the palate (Fig. 80). The commonest case is its deep
bifurcation behind (Fig. 7 9), each fork uniting with the palate bone
of its own side, and sometimes also with the pterygoid. Such is
usually the fixture of the bone behind, and it then rides along as
well as simply bestrides the rostrum. The anterior end of the
vomer may be perfectly free, projecting into the floor of the nasal
chambers (Figs. 62, 77), or the fore end may be variously steadied
or connected with maxillary processes (Fig. 78). When free in
front, and often when not, the vomer is a simple share-like plate,
more or less expanded vertically, quite thin laterally, and “ spiked,”
i.e. running forward to a point; under these circumstances it may
or may not bifurcate behind, and be there attached to the palatines
or not. But the commonest case of vomer, shown by the great
Passerine group, which comprise the majority of recent birds, is
different from this, the vomer being in front thickened, flattened
and expanded laterally, and connected with nasal cartilages and
ossifications (alinasals and turbinals). Such a vomer, deeply cleft
behind to join the palatals, is endlessly diversified in the configura-
tion of its fore end, which may be notched, lobbed, clubbed, etc.
The general case of such a vomer is indicated by the expression
“ vomer truncate in front,” as distinguished from the simply pointed
or “spiked” vomer. (For further details see description of the
several patterns of palatal structure, beyond.)

SEC. IV

ANATOMY OF BIRDS

239

The Quadrate Bone (Lat. guadratus, squared ; Figs. 62, 63, n,
64, 65, 68, 69, 71, q, 75, Qu), with which we may begin the jaw-
bones proper, is the suspensorium of the lower jaw, — the perfectly
constant and characteristic bone by means of which the mandible
proper articulates with the skull. Its rudiment is seen in the
earliest embryos, at the corners of the primordial parachordal
cartilages. It belongs to the mandibular (first postoral) arch, of
which it is the proximal element. Its general morphology has
caused much dispute. From the fact that in birds one of its func-
tions is to support, in part, the tympanum of the ear, it has been
identified with the tympanic bone of mammals, — that which in man
forms the bony tube of the external auditory meatus. The view
now generally accepted is, that the bird’s quadrate represents, cer-
tainly in part, probably in whole, the little bone of the middle ear
called the malleus in mammals. However this may be, the quadrate
of a bird bears the proximal ends of both jaws, carrying their final
(posterior) articulation up to the squamosal and petrosal bones.
Thus, the foot of the quadrate forms the free hinge of the lower
jaw, and also movably articulates the back end of both the zygo-
matic and the pterygopalatine bars or “arcades.” The head of
the quadrate freely articulates with the squamosal, just in front of
the tympanic cavity, which it thus bounds in front ; and there is
usually a shoulder which furthermore articulates with the anterior
periotic bone, the prootic. Struthious birds do not have these two
distinct facets. A long pedicle or orbital process extends forward,
inward, and upward in the orbit ; this non-articular handle is for
advantageous muscular traction. So circumstanced, the quadrate
is a stocky bone, of a shape reminding one of an anvil ; it rocks
freely to and fro upon its cranial socket, pulling and pushing upon
the whole maxillary and mandibular mechanism, with such effect
that when the lower jaw drops, the zygomatic and palatal bars are
automatically shoved forward, tending to make the upper jaw rise,
and so increase the opening of the mouth. Such mobility of the
upper jaw automatically with the movement of the lower is very
free in parrots, whose craniofacial connections are quite articular in
character ; it is well shown also in ducks \ and probably nearly all
birds have some such motion of the upper jaw upon the skull. In
nearly all birds, the mandibular articular facet of the quadrate is
divided by a lengthwise impression into inner and outer protuber-
ances, or condyles, fitting corresponding depressions on the articular
face of the lower jaw in some birds the articular surface is single.
The zygomatic articulation with the quadrate is made by the balled
end of the quadratojugal socketed in a cup at the outer side of the
mandibular facet (with various minor modifications in different
birds). The palatal articulation is made by a little condyle of the

240

GENERAL ORNITHOLOGY

PART II

quadrate, at the inner side of the main facet, socketed into the
cupped end of the pterygoid (with minor modifications).

The Quadpatojugal and Jugal Bones (Lat. jugum, a yoke;
Figs. 62, 63, q, r, 69, 71, qi, j) form most of the outer arcade — the
jugal or zygomatic bar — leading from the quadrate bone to the beak.
The quadrato jugal is posterior, reaching a variable distance for-
ward ; at its fore end it is obliquely sutured to the jugal, a splint-
rod which carries the bar forward to the maxillary bone, with which
it is in like manner obliquely sutured. The whole affair is almost
always a slender rod, which, with its fellow of the opposite side,
forms the outermost lateral boundary of the skull for a great dis-
tance. It corresponds in general with the “ zygomatic arch ” of a
mammal, which is made up of a “ zygomatic process of the squa-
mosal ” and a malar or “ cheek-bone.” The whole zygomatic arch,
including the maxillary bone itself, is developed from the outer part
of the primordial pterygopalatine bar (see Fig. 65). In parrots the
zygoma is movably articulated before as behind.

The Maxillary Bone (Lat. maxilla, upper jaw-bone; Figs. 62,
63, s, 69, 71, 75, mx), forming so much of the upper jaw of a
mammal, is in birds greatly reduced, being starved out by the pre-
dominant premaxillaries which form most of the upper beak. The
shape of this stunted bone varies too much to be concisely described.
Its connections are, ordinarily, with the jugal behind, by a long
slender splint-like process, and with the premaxillary and usually
the nasal bones in front and externally. Internally, it may or may
not connect with the palatal and vomer. The zoological interest of
this bone centres in certain inward (palate-ward) processes, often its
most conspicuous parts, and apparently corresponding to the plate
which in a mammal roofs the hard palate anteriorly. Though these
are mere processes from the main maxillary, they are so distinct
and important that they are commonly described as if they were
independent bones, under the name of the maxillopalatines. They are
flange-like or scroll-like plates, or large spongy masses of delicate
bone-tissue, — endlessly varied in configuration and context (see the
various figures of base of skull, mxp, beyond, where the palatal
patterns are described). Certain other inward maxillary processes,
which may or may not unite with the vomer, and so bridge over
the palate, are called sejjfo-maxillaries (Fig. 80, smx) ; and in some
woodpeckers yet other palatal processes appear (Fig. 80, pmx).

The Pterygoid Bones (Cr. Trrepv^, pterux, wing ; eTSos, eidos,
form; Figs. 62, 63, o, 65, 66, 68, 69, 71, 80, 75 to 79, P/).

Keturning now to the quadrate, and going along the inner arcade,
we first encounter the pterygoid, — a generally rod-like, but variously
twisted, crooked, or expanded bone, which makes the connection
between the quadrate behind and the palate bone before. The

SEC. IV

ANATOMY OF BIRDS

241

pterygoid is always freely jointed at both ends ; its posterior quad-
rate articulation has been noted above ; its anterior connection is
usually by little nipper-like claws by which it “ catches on ” to the
hind end of the palatine. In the ostrich (Fig. 7 5, Pt) the pterygoid
expands into a scroll-like plate ; but its rod-like shape is usually
preserved. Besides passing very obliquely inward as it goes for-
ward from the wide-apart quadrates to the narrow rostrum in the
axis of the skull, the pterygoid often bellies or elbows inwards in
its course to join the basisphenoidal beak, and be movably articu-
lated therewith. In the majority of birds there is no such rostral
articulation, or the pterygoid only touches the rostrum at its fore
end, where it joins the palatal. In many, however, special articu-
lar facets, called hasi;pterygoid ^processes (Fig. 7 0, ap), are developed on
the rostrum for the pterygoids to abut against and glide over. In
Carinate birds, excepting the tinamous {Dromceognathce), these
processes are forward on the beak, and the pterygoids articulate at
or near their own fore ends, as well shown in the fowl or duck
(Figs. 77, 78, Pt). InEatite birds and tinamous, the basipterygoids
are very long, flaring transverse processes, far back on the rostrum,
at the sphenoidal base, and the pterygoids articulate therewith at or
near their own posterior ends (Figs. 7 5, Btp, and 7 6).

The Palatal or Palatine Bones (Lat. palatum, roof of the
mouth; Figs. 62, 63, p, 65, 66, 68, 69, 71, 77, 78, 80, pa, 75,
76, 79, PI) are a pair, approximately parallel and near the mid-line,
forming that part of the “ hard palate ” or roof of the mouth which
is not constructed by the palatal processes of the maxillaries, or by
the vomer. They are nearly always long thin bones, among the
most conspicuous parts when the dried skull is viewed from below.
Sometimes, as in the ostrich (Fig. 15, pi), they are remote from the
axis of the skull and only connected in front with the maxillaries
and maxillopalatines. In many birds they skip the maxillary parts
in going forward to be fused with the premaxillaries ; in most, prob-
ably, they form anterior connections in one or another fashion
with palatal parts both of maxillaries and of premaxillaries.
Behind, they always correctly articulate with the pterygoid. The
mid-line connections made in most Carinate birds (not in Dromceo-
gnathce) are variously with the vomer, with the rostrum, with each
other, or some or all of these relations at once. A long deeply-cleft
vomer may by its posterior forks attach itself to the whole palatal
mid-line, excluding the palatals from the rostrum ; less extensive
attachment of the same kind may permit the palatals to touch each
other and the rostrum posteriorly, while cutting them off anteriorly;
also, a non-cleft vomer may attach itself to the posterior extremity
of the palatals, and bear them off the rostrum. The whole hard
palate may fuse into an indistinguishable mass ; and in almost any
R

242

GENERAL ORNITHOLOGY

PART II

case the relations of the palatals to each other and their connections
afford some of the most valuable zoological characters of great groups
of birds. (Details figured and described beyond.) Though very
variable in configuration, as well as in connections, certain parts of
a palatal may usually be recognised, and conveniently named for
descriptive purposes. Anteriorly, in the great majority of birds, of
whatever technical kind of palatal structure, the palatals are simply
prolonged as flat strap-like or lath-like bars running past the maxil-
lary to the premaxillary region \ and such simple band-like charac-
ter may be preserved behind. Ordinarily, however, the palatals
expand posteriorly, becoming more or less laminar; and in this
plate-like part three surfaces may usually be recognised. One, more
or less horizontal, flaring outward, is the external lamina. It is well
shown in a Passerine or Eaptorial bird, where the 'podero-external
angle (between the outer border and the posterior end) of the pala-
tal is well marked, or may be acutely produced ; there is no such
lamina in a fowl, where the palatals are for the most part slender
and rod-like. An internal plate, more or less vertically produced
to make the mid-line rostral or vomerine connection, is the superior
internal lamina^ or mediopalatine process ; very strong, for example,
in a fowl, where it forms all the expanded part of the bone, and
ends anteriorly as a sharp inter-palatine spur. The mediopalatine
is probably to be regarded as the main body of the bone, being the
most axial part, of the most extensive and varied connections. A
third lip or plate of the palatal is the inferior internal lamina, look-
ing downward ; it is generally very evident, but in a duck or fowl
is reduced to a mere ridge, indicating where the superior internal
and external laminae meet. A duck’s palatals are quite different in
appearance from those of most birds, all the posterior parts just dis-
tinguished been reduced and constricted, while the fore ends,
running abruptly into the hard-boned beak, are much expanded
horizontally (Fig. 78). The postero-external angles of the palatal
(formed by the external lamina), even when much produced, may
not reach as far back as opposite the pterygopalatine articulation ;
or they may surpass these limits, and when they do, such backward
prolongation is called postpalatine, the palate being considered to
end at the pterygoids. In like manner, the maxillary processes of
the palatals, or the palatal strips as prolonged into the premaxillary
region, are called prepalatines. The inner posterior process, by
which the palatine is articulated with the pterygoid, is its pterygoid
process.

The Premaxillary Bones (Figs. 62, 63, 69, 70, 71, 80, px,

75 to 7 9, pmx), also called Intermaxillaries, form most of the upper
beak, attaining enormous development in birds, and reversing the
usual relative size of premaxillary and maxillary. Mainly deter-

SEC. IV

ANATOMY OF BIRDS

243

mining as they do the form of the upper mandible, their shapes are
as various as the bills themselves of birds ; but their generalised
characters can be easily given. Each premaxillary, right and left,
forms its half the bill; the two are always completely fused
together in front, commonly preserving traces at least of their
original distinction behind. They are commonly called one bone, the
premaxillary. Each is a triradiate or 3-pronged bone ; one upper
prong, the most distinct, called the nasal or frontal process, forms
with its fellow the culmen (Fig. 26, h) of the bill. These pro-
cesses, side by side, run clear up to the frontal bone in birds,
driving the nasal bones apart from each other. Such a median
fronto-premaxillary suture, with lateral frontonasal and nasopre-
maxillary sutures, is highly characteristic of birds, — an arrangement
probably exceptionless. Two other horizontal prongs on each side,
extensively distinct from the frontal process in most birds, but
less separate from each other, run horizontally along the side and
roof of the mouth for a variable distance. These horizontal prongs
are an external or dentary process (Fig. ^0,ppx), forming the tomium
(Fig. 26) of the bill, and reaching back to join the dentary part of
the maxillary; and an internal or palatal process (Fig. d>0, ppx), run-
ning along the commencement of the bony palate. With this latter
the anterior ends of the palatal bones unite, — either on the side to-
ward the mid-line of the beak, or between the palatal and dentary
processes, as in a woodpecker (Fig. 80). Great laminar expansions
inward of these palatal parts of the premaxillaries roof the hard
part of the mouth anteriorly, though there is usually a vacancy
between the premaxillary hard palate and that formed farther back
by the maxillopalatines and palatines. The posterior extremities
at least of the frontal processes of the premaxillaries are commonly
distinguishable from each other as well as from the frontal and
nasal bones — in fact, these fronto-naso-premaxillary sutures are
among the most persistent of all. The divergence of the frontal
from the palatal and dentary processes bounds the external nostril
in part, the circumscription of that orifice being completed by the
prongs of the nasal bones. The superficies of the premaxillary
bone, like that of the dentary piece of the lower jaw-bone, is com-
monly sculptured with the impressions of the vessels and nerves
which ramify beneath the horny integument ; and in birds with
very sensitive bills, as a snipe or duck, the end is perforated sieve-
like with little holes, into which the skin shrinks in drying, produc-
ing the familiar pitted appearance (Fig. 63, at c).

The Nasal Bones (Figs. 62, 71, ?^) might have been described
next after the frontals, as they continue forward the general roofing
of the skull ; but are conveniently considered in the present con-
nection, being in birds rather “facial” than “cranial.” They are of

244

GENERAL ORNITHOLOGY

PART II

large size, and pronged, — one fork, the superior process, being applied
for a variable distance along the outer side of the frontal process of
the premaxillary, the other, inferior, descending to or towards the
dentary border of the maxillary or premaxillary, or both ; the
divergence of these two processes bounding the nostril behind. The
base of the nasal, uppermost and posterior, ankyloses (usually) or
sutures (often) or articulates (as in parrots) with the antero-external
border of the frontal bone ; its frequent collateral connections being
with the lacrymal or ethmoid, or both of these. The nasals are
very variable in shape, as well as in the extent of their connections.
When expansive, they may wall in much of the nasal cavity, as
well as bound the nostrils. These latter openings, as far as their
bony boundaries are concerned, are usually much more extensive
than they seem to be from the outside, being greatly contracted by
membrane and integument. Ordinarily, each forms a large vacuity,
which the descending prong of the nasal bone separates from a
similar vacancy between itself and the lacrymal, the lacrymal in turn
interposing between this and the orbital cavity. The descending
process of the nasal, in fact, is a marked object at the side of the
base of the upper mandible of most birds, though slight or rudi-
mentary in the Eatitse. A character of the nasals has been
employed in classification by Mr. Garrod. A bird having the
bones as above generally described, with moderate forking, so that
the angle of the fork, bounding the nostrils behind, does not reach
so far back as the fronto-premaxillary suture, is termed holorhinal
(Gr. oAo?, holos, whole j pt?, ptvo?, rhis, rhinos, nose ; Fig. 62). But
in the Columhidce, and in a great many wading and swimming
birds, whose palates are cleft {schizognathous), the nasal bones are
schizorhinal (crxiC^, schizo, I cut) ; that is, cleft to or beyond the
ends of the premaxillaries ; such fission leaving the external de-
scending process very distinct from the other, almost like a separate
bone. Pigeons, gulls, plovers, cranes, auks, and other birds are
thus split-nosed. The value of the character, except as an auxiliary,
is doubtful.

The Lacrymal (Lat. lacryma, a tear; from the relation of the
human bone to the tear-duct; Figs. 62, 63, u, 71, 1) is one of
several splint-like membrane-bones of the skull, having little inti-
macy of relation with the general morphology of the cranium,
though quite constant in birds, and often very conspicuous. It is
situated at or near the anterior outer corner of the orbit, near the
nasal but behind that bone ; sometimes ankylosed, sometimes very
loosely attached, oftener firmly sutured with the frontal ; and may
also have connection with the nasal and ethmoid. It is generally a
claw-like affair, depending from the front outer corner of the frontal,
and consequently bounding the orbit anteriorly ; it may be variously

SEC. IV

ANATOMY OF BIRDS

245

twisted, crooked, hooked, etc. It is singularly elongated and dis-
torted in the ostrich. In the duck tribe, in which the lacrymo-
frontal region of the skull is greatly elongated, the lacrymal has
coextensive attachment to the frontal bone, and is broadly laminar,
with a downward process ; in some ducks bounding at least a fourth
of the orbital brim, and almost completing the circle by extending
toward the very protrusive postfrontal process, as in Fig. 63, u. In
some parrots, the rim of the orbit is completed below, and even
sends a bony bar to bridge over the temporal fossa behind the
postfrontal. In some birds, the lacrymal is quite free, and even
in more than one free piece. The os uncinatum, or os lacrymoFOila-
tinum, would appear to be a palatine bone distinct from the lacrymal ;
it has been observed in the Musophagidce and many other picarian
birds, in Tachypetes, and certain ProcellariidcB. The lacrymal bone
seems to be the principal relic, in birds, of a set of splint-bones
which lie about the edges of the orbits in many Sauropsida. Another
is the postfrontal or sphenotic, usually a process of the frontal,
often a separate ossification. In some birds, as various Piaptores,
there are one or more loose supraorbital plates of bone, serving to
eke out the brim of the orbits ; thus forming the “ orbital shields ”
so prominent in many hawks, and causing their eyebrows to pro-
ject. Were such a chain of splint-bones complete (lacrymal,
superorbitals, postfrontal, and squamosal, to quadrate), it would
form an arcade of bones, over the orbit, like the actual zygomatic
arch (maxillary, jugal, quadratojugal, to quadrate) which lies under
the orbit ; and such a double series is very perfectly illustrated in
many of the Sauropsida below birds.

Other special ossifications have been described in some birds,
but I am obliged to pass them over. I have already far exceeded
intended limits, and have yet to describe the mandibular and
hyoidean arches, and the zoological characters of the palate as a
whole.

The Mandible, op Lower Jaw-Bone (Figs. 62, 63, 70, 71) is a
collection of bones developed in the first postoral visceral arch.
Each half of the compound bone (right and left) consists normally
of jive bones, which become immovably ankylosed, but traces of the
original distinction of which commonly persist for an indefinite
period, — in some birds throughout their lives. In an embryo whose
skull has passed to the cartilaginous stage, a long slender rod of
cartilage appears in the first postoral visceral arch ; this is MeckeVs
cartilage, or the MecTcelian rod (Figs. 65, 66, 68, 70, mk), so named
after a famous anatomist. Around this rod, which subsequently
disappears, the several bones of the mandible are developed. The
anterior one of these is the dentary (d), forming the scaffold of the
horny part of the external under mandible. It usually unites by

246

GENERAL ORNITHOLOGY

PART II

ankylosis, sometimes only by suture, with its fellow of the opposite
side. This union in the middle line is the symphysis (Gr. o-vv, sun,
with ; (f>v(TL^, phusis, growth). The line of union is externally the
gonys (see antea), the length and other characters of which are
determined by the mode of symphysis, as is the general shape of the
tip of the lower mandible. The union generally makes an angular
A, but may be an obtuse fl the symphysis is very short and im-
perfect, as in a pelican, for instance, or the opposite, as in a wood-
pecker and a multitude of birds. Behind the dentary, each ramus of the
jaw continues with pieces called splenial, angular and surangular {sp, a,
su) y there is often a fenestra between them, by imperfection of bony
union, as shown in Fig. 62 or 63,/, which also sufficiently indicates
the relations of these parts. The articulation of the jaw with the
quadrate bone is furnished by a fifth piece called articular (ar) from
its function. As a whole the mandible is a pronged bone, forking
with a variable degree of divergence from its obtuse or acute point,
sometimes quite parallel-sided, as in a duck, oftener very open ;
such prongs may be straight, or variously curved or bent either in
the vertical or the horizontal plane ; are generally stout and stanch,
sometimes so slender as to be quite flexible. The articular part,
always expanded horizontally, presents a smooth irregularly cupped
superior surface for reception of the protuberances of the foot of
quadrate. In general, this concave articular surface is divided into
an inner and outer cup separated by a protuberance, corresponding
to similar inequalities of the opposing surface of the quadrate.
Cupping of the mandibular articulation is characteristic of birds
as compared with mammals, in which latter the lower jaw has
always a knobbed articular surface (condyle). In many birds
the angle of the jaw is prolonged back of the articulation as a
posterior articular process (Fig. 63, A, 70, 71, pap), which may be long,
slender, and upcurved, as is well shown in a fowl, duck, or plover.
Such birds are said to have the “ angle of the mandible recurved ; ”
the opposite condition is “ angle truncated ” (cut off). Usually also,
an internal angular process (Figs. 70, 71, iap) is produced inward from
the articular part of the jaw, as in the fowl or duck. Between the
dentary and articular parts, the ramus of the jaw is usually vertically
produced as a thin raised crest, which, when prominent, is called
the coronoid process ; it corresponds to the strong process so called in
a mammal, and relates to the advantageous insertion of the temporal
or masseteric muscles which effect closure of the jaw. It is scarcely
evident in the fowl (Fig. 62), but well marked in the duck (Fig. 63,
over /). At the back of the articular surface is the pneumatic
foramen for entrance of air, when any ; on the inner surface of the
ramus, about the splenial bone, is the opening conveying the vessels
and nerve.

SEC. IV

ANATOMY OF BIRDS

247

The Hyoid Bone (Gr. letter v, hu = hy, etSo?, eidos, form ; Figs.
65-68, 72-74) is the skeleton of the tongue ;
a very composite structure, consisting of
several distinct bones, developed in the
second and third postoral visceral arches (see
Fig. 65, where ch and Wi are the original ele-
ments of the second arch, making the hasi-
hyal ^ and ceratohyal bones, and Ihr, cbr, and
ehr, are the original elements of the third
arch, making the basibranchial, ceratobranchial,
and epibranchial bones). The whole affair
is somewhat A“ or f| -shaped, lying loosely,
point forward, between the forks of the
lower jaw, with its long slender prongs curv-
ing up behind the hind head more or less ;
but it is not definitely connected with any
other bones of the skull. The connection
which exists between the hyoid and other
cranial bones in a mammal is in birds broken
by non-development of certain links of bone
developed in the mammalian second post-
oral arch, as the stylohyal, epihyal, etc. ;
though birds have a rudimentary stylohyal,
at least in the embryo, among the several
proximal parts of the second arch which form
the intricate bones within the ear-passages
(Fig. 67). The visible parts of a bird’s hyoid
are usually : the body of the bone, basihyal
(bh, and Fig. 72, c), single and median, com-
monly quite short and stocky, sometimes long
and slender. The basihyal bears in front a
pair of ceratohyals (ch ; not shown in Fig. 7 2,
where they have been absorbed in b) usually
movably articulated with the basihyal. They
commonly appear as little “ horns ” or pro-
cesses of the next piece, the glossohyal (Fig.

72, b) or bone chiefly supporting the sub-
stance of the tongue. It may be a stout and
apparently single bone, as that of the goose jiyaisor “fesse^ cornua’’; c,

n 1 t , o, . ~ basihyal, movably articulated

figured ; but oitener appears as a pair 01 with &, and combined com-
slender bones, side by side, whose backward cafied‘‘‘’uSy^^^^

ends are the ceratohyals._ The glossohyal i

may or may not bear at its fore end a car- known as “ thyrohyais,” or
tilaginous tip, as in Fig. 72, a. All the greater cornua.

^ Basihyal, etc. The word hyal, used only in composition, means the same as

Fig. 72.— Hyoid bones of a
goose, nat. size ; by Dr. R. W.
Shufeldt, U.S.A. a, carti-
laginous end-piece of h, the
great glossohyal, which has
absorbed or replaced cerato-

248

GENERAL ORNITHOLOGY

PART II

foregoing are hyal, i.e. belonging to the second visceral arch ; the
following are branchial, of the third arch : The hasihranchial {bbr ;
Fig. 72, d) is a single median piece, projecting backward from the
basihyal, with which it may be perfectly consolidated, as it is in the
figure, or separately articulated ; it may be wanting ; it is usually
tipped and prolonged backward with a thread of cartilage. The
hasihranchial is oftener called “ urohyal,” but had better be allowed
its strict morphological name. On either side, the basihj^al bears
the separately articulated ceratobranchials {cbr; Fig. 72, e), long
slender bones diverging as they pass backward, and bearing upon
their ends the epibranchials (ebr; Fig. 72, /), which finish off the

Figs. 73, 74. — Under Fig. side view of a woodpecker’s (Picus) skull, showing the long
slender basihyal (hh), bearing slight elements at its fore end, no uroyhal, and extraordinarily
long thyrohyals (c6r, ehr) curving up over back of skull and curling together around orbit
of the right eye. Upper Fig. top view of skull of Colaptes, showing thyrohyals running along
the skull and into right nostril to end of the bill. (Dr. R. W. Shufeldt, U.S.A.)

hyoid bone behind, or may be in turn tipped with cartilaginous
threads. The cerato- and epibranchials together are badly called
the “ thyrohyals,” and in still more popular language the “ greater
cornua ” or “ horns ” of the hyoid. All these bones vary in differ-
ent birds in size and shape and relative development ; the branchial
elements are the most constant in their length and slenderness.
The whole hyoid apparatus of the woodpeckers is specially modi-
fied ; the basihyal is very long and slender, bearing stunted cerato-
and glossohyals at its extreme end ; there is no urohyal, or only

hyoid, t)ut it is said of the several different elements of which the hyoid hone, or
hyoidean arch, is composed, the termination -al being conformable with hrancM-al,
etc.

SEC. IV

ANA TOMY OF BIRDS

249

a rudiment ; the ceratobranchials are long, and the epibranchials so
extraordinarily elongated in some species as to curl up over the back
of the skull and forward along the top of the skull to a variable
distance ; sometimes, as in Fig. 7 3, curling around the orbit of the
eye, or, as in Fig. 74, running into the nostril to. the tip of the beak.
In such cases they bundle together in passing forward over the
skull, and go obliquely to one side.

Other Bones of the Skull. — The articulation of the lower jaw
with the quadrate may have certain sesamoids. Thus, there are two
such sclerosteous or ligament-bones in the external lateral ligament
of the raven’s jaw-joint, and the long occipital style of the cor-
morant and snake-bird is of the same character, being an ossification
in the nuchal ligament of the neck. The siphon-like tube which
conveys air from the outer ear-passage to the hollow of the mandible
may ossify, as it does in an old raven, for example, resulting in a
neat tubular “ air-bone ” or atmosteon (Gr. ar/^os, air).

Types of Palatal Structure. — The arrangement of the bones of
the palate in birds results in several types of structure, first defined
by Huxley and applied to the classification of birds. These are the
dromceognathous^ schizognathous^ desmognathous^ and mgitJiognathous ; to
which Parker has added the saurognathous. Huxley proposed to
make the primary division of Carinate birds upon this score ; and
since the plan could not be made to work in his hands, it is certainly
futile for any one else to demonstrate again the impossibility of
establishing the higher groups of birds upon any one set of charac-
ters,— upon the modifications of any one structure. Nevertheless,
when duly co-ordinated with other characters, palatal structure
becomes of the utmost importance in defining large groups of birds.
It is necessary, therefore, for the student to clearly understand this
matter, which I will lay before him as nearly as possible in the words
of the authors just mentioned.

Dromseognathism (Gr. Spofialos, dromaios, a runner : genus-name
of the emeu). All the Eatite birds, and the tinamous alone of
Carinate birds, are dromcBognathous. “ The posterior ends of the
palatines and the anterior ends of the pterygoids are very imperfectly,
or not at all, articulated with the basisphenoidal rostrum, being
usually separated from it, and supported by the broad, cleft, hinder
end of the vomer. Strong basipterygoid processes, arising from the
body of the basisphenoid, and not from the rostrum, articulate with
facets which are situated nearer the posterior than the anterior ends
of the inner edges of the pterygoid bones.” This is the gist of
dromoeognathism ; it is exhibited in several ways, (a) In Struthio
alone. Fig. 75, the very short vomer, borne upon the rostrum, articu-
lates neither with palatines nor with pterygoids, but with the maxillo-
palatines ; and the palatines, which are remote from the rostrum,

SEC. IV

ANATOMY OF BIRDS

251

advance beyond the maxillopalatines, as in most birds. (5) In Rhea,
the vomer is as long as usual in birds, and articulates behind with
the palatines and pterygoids, but does not join the maxillopalatines
in front ; the short palatines unite with the inner and posterior
edges of the thin fenestrated maxillopalatines. (c) In Casmrius
and Dromoeus (cassowary and emeu) the long vomer articulates
behind with the palatines and pterygoids, and unites in front with
the maxillopalatines j these are flat, imperforate, and solidly joined
to the premaxillse ; the palatines are short, (d) The extinct Dinornis

Fig. 76. — Dromceognathous
skull of tinamou (Tinamus ro-
hustus) ; copied by Shufeldt from
Huxley. Letters as before.

Fig. 77. — ScMzognathous skull of com-
mon fowl, nat. size ; from nature by Dr. R.
W. Sbufeldt, U.S.A. Letters as before,
except Pa, palatine.

had flat imperforate maxillopalatine plates uniting solidly with the
premaxillse, and probably with the vomer, as in Dromceus. (e) In
Ajpteryx the long vomer unites with palatines and pterygoids behind ;
short broad palatines suture obliquely with flat imperforate maxillo-
palatine plates, which unite both with premaxillary and vomer. (/)
The tinamous, Dromceognathce (Fig. 76) “have a completely struthious
palate ” ; vomer very broad, uniting in front with broad maxillo-
palatine plates as in Drommus ; behind articulating with posterior
ends of palatines and anterior ends of pterygoids, both of which are
thus prevented, as in all Ratitce, from any extensive connection with
the rostrum ; basipterygoid processes springing from body of

252

GENERAL ORNITHOLOGY

PART II

sphenoid, not from its rostrum, articulating with pterygoids very
near the posterior or outer ends of the latter ; and head of quadrate
with a single articular facet, as in Batitce.

Sehizognathism (Gr. o-xtC(a, schizo, I cleave) is the kind of
“ cleft palate ” shown by the columbine and gallinaceous birds, by
the waders at large, and many of the swimmers (see' Fig. 77). In
this general case, the vomer, whether large or small, tapers to a
point in front, while behind it embraces the basisphenoidal rostrum,
between the palatines ; these bones and the pterygoids are directly
articulated with one another and with the basisphenoidal rostrum,
not being borne upon the divergent posterior ends of the vomer ;
the maxillopalatines, usually elongated and lamellar, pass inwards
over [under, when the skull is viewed upside-down, as it usually is]
the anterior part of the palatines, with which they unite and then
bend backwards, along the inner edge of the palatines, leaving a
broader or narrower fissure between themselves and the vomer, on
each side, and do not unite with one another or with the vomer.
It follows from this that in the dry skull of a plover, for instance,
which shows the schizognathous arrangement extremely well, “ the
blade of a thin knife can be passed, without meeting with any
bony obstacle, from the posterior nares alongside the vomer to the end
of the beak.” There are several groups of birds which exhibit the
schizognathous plan, with ulterior modifications of palatal and other
characters, (a) The columbine birds (Feristeromorphce of Huxley’s
arrangement) : maxillopalatines elongate and spongy ; basiptery-
goid processes narrow, but prominent, (b) The gallinaceous birds
(Alectoromorphce) : maxillopalatines varying greatly in size, but always
lamellar : palatines long and narrow, with rounded -off postero-
external angles ; basipterygoid processes oval, flattened, sessile upon
the rostrum, articulating with the pterygoids, (c) The penguins
{Spheniscomorphce) : maxillopalatines concavo-convex and lamellar ;
no basipterygoid processes ; pterygoids flattened, (d) In the gulls,
petrels, loons, grebes, and auks, constituting the Cecomorplice, the
maxillopalatines are usually lamellar and concavo-convex, but may
be spongy, tumid, and closely approximated to the vomer ; and
basipterygoid processes are absent or present, {e) In the cranes,
rails, and their allies (Geranomorpli(E), the maxillopalatines are
concavo-convex and lamellar, and basipterygoid processes are usually
absent. (/) In the plover- snipe group, or limicoline Grallce
(Charadriomorphce), the maxillopalatines are always concavo-convex
and lamellar ; the basipterygoid processes narrow and prominent.
Excepting perhaps group d, which does not hang together so well,
the schizognathous groups here noted correspond very closely with
recognised orders or suborders of birds ; in all of them, the maxillo-
palatines are perfectly distinct from one another and from the vomer.

SEC. IV

ANATOMY OF BIRDS

253

and the latter is slender and usually pointed. There are plenty of
other birds in which the former factor in the case obtains ) but in
these the vomer is broad and usually truncate in front (see u^githo-
gnathism, beyond).

To.,

Desmognathism (Gr. Sea-fios,
desmos, a bond) is exhibited in
one or another style by those
swimming and wading birds
which are not schizognathous,
by the birds of prey, and
various non-passerine perching
birds. It does not fadge so
well as any other one of the
palatal types of structure with
recognised groups of birds
based on other considerations.

In this “ bound -palate ” type
the vomer is either abortive or
so small that it disappears ;
when existing it is usually
slender and tapers to a point
in front; the maxillopalatines
are united across the median
line, either directly or by means
of ossifications in the nasal sep-
tum ; the posterior ends of the
palatines and the anterior ends
of the pterygoids articulate
directly with the rostrum (as in
schizognathism). This type is
simply and perfectly exhibited
by a duck (Fig. 78) in which
the maxillopalatine is a broad
flat plate united with its fellow
in mid-line ; the oval sessile
basipterygoid facets are far for-
ward, opposite the very ends
of the pterygoids. In the
flamingo, ibis, spoon-bill, stork,
heron, the united maxillo-
palatines are tumid and spongy, Ailing the base of the beak ;
basipterygoids are wanting (rudimentary in the flamingo). In
totipalmate swimmers (pelican, cormorant), desmognathism is carried
to an extreme by union of the palate bones also across the mid-line ;
the general arrangement is as before. The birds of prey exhibit

Fig, 78. — DesmognatJious skull of mallard
duck, Anas hoscas, nat. size ; from nature by Dr.
R. W. Shufeldt, U.S.A. Letters as before.

254

GENERAL ORNITHOLOGY

PART II

several special conditions of desmognathism. The parrots are
another case ; among other cranial characters of these birds is to be
noted the articulation of the palate bones with the upper beak, like

that of the zygoma. The multi-
farious Picarian birds, or non-
passerine Insessores, are desmo-
gnathous, excepting the schizo-
gnathous trogons (Trogonidce) and
the “saurognathous’’ woodpeckers.
Parker has described the follow-
ing categories of desmognathism :
(a) Perfect direct, the maxillopala-
tines uniting below at the mid-
line; either with the nasal septum
free from such bony bridge, as in
a duck; or ankylosed therewith,
as in many birds of prey, ih)
Perfect indirect, very common, as
in eagles, vultures, owls ; maxillo-
palatines separated from each
other by a chink, but ankylosed
with nasal septum, (c) Imper-
fectly direct ; maxillopalatines
sutured together but not anky-
losed. “In young falcons and
hawks the palate is at first in-
direct, is then imperfectly direct,
and at last perfectly direct.” (d)
Imperfectly indirect; maxillopala-
tines closely articulated with, and
separated by, the “ median septo-
maxillary ” ; but there is no
ankylosis, {e) Double: the pala-
tines united as well as the maxillo-
palatines ; as in the pelican and
cormorant above noted, in certain
Caprimulgine birds, horn-bills, etc.

J.11C xxxxc, »- , LU . 7 1,1 1

the ossified nasal septum borne upon the end (/) Compound .* when the properly
of the vomer, which latter bone begins at the •ii i n r

thickest part of the central projection. Mxp CegithognathoUS skull 01 a paSSer-
nnderlies V and overlies PI, but touches beCOmeS also desmO-

neither,

gnathous.

.^githognathism (Gr. alyiOaXos, aigithalos, some small bird) is
exhibited almost unexceptionally by the great group of Passerine
birds ; it is also nearly coincident with Passeres, though a few other
birds, notably the swifts (Cypselidee), also exhibit it. Huxley’s term

Fig. 79. — ^githognathous skull of raven,
Corvus corax, nat. size ; from nature by Dr. R.
W. Shufeldt, U.S.A. Letters as before.
N.B. The misplaced reference line, V, goes to

SEC. IV

ANATOMY OF BIFFS

255

Gm'acomorphcBj nearly synonymous with Passeres, relates to the
palatal structure exhibited by a raven (Fig. 79), as typical of that of
Passeres at large. The vomer is a broad bone, truncate in front and
deeply cleft behind, embracing the sphenoidal rostrum in its forks.
The palatines have produced postero-external angles. The maxillo-
palatines are slender at their origin, extending inward and back-
ward over the palatines and under the vomer, where they end free,
being united neither with each other nor with the vomer. This
disconnection of the maxillopalatines is ,quoad hoc “ schizognathous,”
of course ; but such condition, in association with the peculiarities of
the vomer, is segithognathous. The nasal septum in front of the
vomer is often ossified in segithognathism, and the interval between
it and the premaxillse filled up with spongy bone; but no union
takes place between this ossification and the vomer (Huxley).
According to Parker, the distinguishing character of the aegitho-
gnathous type is the union of the vomer with the alinasal wall and
turbinals. He distinguishes four styles ; (a) Incomplete ; very curi-
ously exhibited by the low Turnix, which stands near the gallina-
ceous birds, (h, c) Complete^ as represented under two varieties,
one typified by the crow, an Oscine Passerine, the other by the
Olamatorial Passerines Pachyrhamphus and Pipra. (d) Compound,
i.e. mixed with a kind of desmognathism, as noted above. “Vomer
truncated in front ” is the general expression for the condition of
that bone in the aegithognathous type ; it is frequently massive in
that direction, and of endlessly varied configuration.

Saupognathism (Gr. a-avpos, sauros, a lizard; Fig. 80). Accord-
ing to Huxley the woodpeckers exhibit a “ degradation and simpli-
fication of the aegithognathous structure.” The peculiarities of the
palate of these birds (including Picidce, Picumnidce, and lyngidce) are
so decided that Parker proposes to call them saurognathous. The
structure is very difiicult to make out, and may be understood best
by study of the accompanying figure, copied from Parker. The
maxillopalatines, mxp, are very slight, not extending inward beyond
the outer margin of the palatines, and being sometimes quite rudi-
mentary. In front of them, an additional little palatal plate of the
maxillary, pmx, is developed. The vomers, v, are delicate paired
rods on each side of the median line. The postero-external angle of
the palatine is either rounded off or obtuse-angled. Where the
broad main part of the palatine suddenly narrows is developed an
interpalatine process, ipa. The ethmopalatine plates, epa, or internal
superior plates of the palatine, which are of variable length, are con-
nected by the most marked mediopalatine ossification, mpa, seen in
the class of birds. Bridges of bone are deposited along the inner
borders of the palatines ; such are the septomaxillaries, smx, and
other formations which, like the mediopalatine, serve to bind the

256

GENERAL ORNITHOLOGY

PART II

palate halves together. The nasal chambers are unusually simple ;
there are peculiarities of the tympanic cavity and quadrate hone.

“ All these things being considered,” says Parker, in conclu-
sion, “it will seem contradictory now to assert the great uniformity

of the skulls of Birds,
and indeed of Birds
themselves. Yet so it
is, and the countless
modifications that offer
themselves for observa-
tion are gentle in the
extreme. One form is
often seen to pass into
another by almost in
sensible gradations. . . .
In the rest of the
Birds’ organisation
abundant evidence of
the same specialisation
will be seen. The mind
fails to desire more
beauty or to contem-
plate more exquisite
adaptations. An almost
infinite variety of Ver-
tebrate life is to be
found in this class. Of
its members some dig
and bury their germs,
which rise again in
full plumage, whilst
others watch and in-
cessantly feed their
tender brood in the
shady covert or ‘ on
the crags of the rock

Fig. 80. — Saurognafhous skull of nestling Pwms miwor, x 4
diameters, after Parker. Px, premaxillary ; dpx, its dentary
process ; ppx, its palatal process ; sn, septonasal ; pa, pala-
tine; pmx, peculiar palatal plate of maxillary of a wood-
pecker ; nf, nasal turbinal ; mx, maxillary ; ipa, interpalatal
spur of palatine bone ; mxp, rudimentary maxillopalatine,

scarcely reaching palatine ; smx, septomaxillary, in several j ^ >

pieces ; v, right vomer, its fellow opposite ; pe, lower border and the Strong place,
of perpendicular plate of ethmoid, between vomers; epa, j InnnTnnti’rkTi
ethmoidal (inner) plate of palatine; mpa, mediopalatine ; lUCUmULlun

pg, pterygoid ; i, foramen for internal carotid ; 8, for vagus -walk OtherS

nerve ; 9, for hypoglossal nerve.

some
run, or
they may wade, swim,
plunge, or dive, whilst most of them ‘ fly in the open firmament of
heaven.’” {Ency. Brit. 9th ed. Art. “Birds,” p. 717.)

SEC. IV

ANATOMY OF BIRDS

257

h. Neurology; The Nervous System; Organs of Special
Senses

The Nervous System of any Vertebrate determines the form of
such an animal ; in fact, the beautiful skeleton we have examined
is simply a sketch in bone of the cerelrospinal nervous system, con-
formably with which the whole bony framework of the body is
erected. A brain and spinal cord and their lateral prolongations
or nerves are the commanding superadditions, in a vertebrate, to
any such nervous system as an invertebrate may or does possess.
Besides the vertebral or main nervous system, all brainy vertebrates
retain a sympathetic system of nerves, supposed to represent a modified
inheritance of the corresponding nervous system of Invertebrates.
Thus the cerebrospinal and sympathetic are the two distinct nervous
systems of all vertebrates which have a skull and brain. The
former presides over the animal life of the creature, — its sensations,
perceptions, and voluntary actions ; the latter more especially over
its vegetative functions, as digestion, respiration, circulation, and
reproduction, which are more or less involuntary. But the two are
inseparably connected, anatomically and physiologically, so that no
distinct line can be drawn between them. Nerve-tissue consists of
an aggregation of nerve-cells and their investing substance, — the
bodies of a myriad Neuramoebce agglutinated by their secretions.
They are of two species : Neuramoeba cinerea and V. Candida. The
former are usually multiradiate, inosculating cells of nerve-substance,
which form the “ gray matter ” of the brain and spinal cord and
the ganglia (knots) of nerves; the latter are white, thready, and
form the connections of the ganglionic masses and the whole sub-
stance of ordinary nerve-cords. The gray amoebas are the imme-
diate communicants between the mind and the body of the creature ;
the white amoebas are the mediators between the body and outward
things. The gray amoebas translate thought in terms of matter,
and conversely; the white convey the translation. How this is
done, no one knows, but the fact is manifest. In ordinary language,
gray nerve centres receive from white tracts impressions made upon
the periphery of the nervous system ; and, with or without the
knowledge and consent of the animal, convert these impressions
into appropriately responsive actions. This is called the “reflex
action " of the nervous system. Some think such reflection is the
principal or only activity of the nerve-tissue, taking animals to be
mere automata, the mechanism of which is only set in motion by
external stimulation. Others think that animals, and even human
beings, have in their consciousness an inner spring of action, vaguely
called “spiritual,” whose operations upon the matter of their bodies

258

GENERAL ORNITHOLOGY

PART II

manifests what is called by some “ mind,” by others “ soul.” I am
satisfied of the correctness, in the main, of the latter view ; but,
however this may be, it is quite certain that white nerve-tissue is a
means of carrying something to and fro, which something is called
a “ nerve impulse,” for want of knowing what it is. White nerves
have therefore an efferent function, when they carry impulses out-
ward from gray centres, and an afferent function, when they bring
impulses into gray centres. The former is their motor function ; the
latter is their sensory function. In nerves at large, impulses of both
kinds travel in the same tracts without interference ; such mixed
nerves are therefore called sensorimotor. Thus, each spinal nerve
has a posterior sensory ganglionated root, and an anterior motor
simple root, which soon blend in one cord, in which both functions
coexist. Some nerves seem to be entirely motor, as those which
move muscles of the face and tongue. The purest sensory nerves
are those of “ special sense,” as the olfactory, optic, and auditory.
Some nerves are so “mixed” as to combine functions of special
sense, common sensation, and motion, as that called glossopharyn-
geal, which moves, feels, and tastes. The motor effluence of nerve-
tissue upon itself and other parts of the body is literally animation ;
the sensory influence is nominally materialisation. The physical
mechanism of these occult processes in a bird is as follows : —

The Brain (Lat. cerebrum ; Gr. iyKecfiaXov, eghephalon, the enceph-
alon) is the anterior dilatation and complication of the main nervous
axis of the body, contained within the skull. It resembles a soap-
bubble blown at the end of a pipe, being not less beautiful in its
iris-quality, and not less lasting. It is primarily triune, or three-
fold, beginning as three such bubbles, called the anterior, middle, and
posterior cerebral vesicles, corresponding to what are afterward the
fore-brain, mid-brain, and hind-brain, or prosencephalon, mesencephalon,
and opisthencephalon. Two of these primitive vesicles subdivide each
into two, so that five segments of the brain result. These five are
commonly called prosencephalon, diencephalon, mesencephalon, epen-
cephalon, and metencephalon, named from before backward ; and they
respectively correspond with parts of the adult brain known as
cerebruni proper, optic thalami, optic lobes, cerebellum, and medulla
oblongata. The birth and multiplication of gray neuramoebas cause
thickenings of the bladdery membranes in various places and ways ;
all such gray deposits are the ganglia of the brain, and the great
peripheral ganglion is the cortical layer or “bark of the brain.”
Similar deposits of white neuramoebas connect all these ganglionic
colonies, furnishing various commissures of the brain. The cavity
of the original bubbles, continuous with the hollow of the pipe-stem
or spinal cord (which was at the outset a furrow along the back of
the embryo, not a tube) becomes partially divided up into several

SEC. IV

ANATOMY OF BIRDS

259

communicating hollows ; these are the ventricles (little bellies) of the
brain. Actual prolongations of brain-tissue, or nervous threads like
the ordinary spinal nerves, pass out of the brain-box; these are
cerebral nerves^ oftener called cranial nerves ; there are twelve pairs
of them. At the pituitary space, just behind which the noto-
chord ends (see Fig. 64), is developed a remarkable structure, the
pituitary body : its nature is unknown. This lies under the brain ;
opposite it, on top of the brain, is another curiosity, the conarium or
'pineal body ; it has been considered the special seat of the soul by
some, though others have located that throne of animal grace in the
solar plexus of the sympathetic system, which is in the belly. It
is probably the remains of an extinct eye. The pituitary and
pineal are also called respectively the hypophysis and epiphysis
cerebri. They lie respectively at the bottom and top of one of the
cavities of the brain, arbitrarily called the third ventricle; the
anterior wall of this ventricle is the lamina terminalis, or terminal
sheet of the brain, with which, morphologically speaking, the brain
ends in front ; though, in its actual growth, the prosencephalon
crowds ahead of this formation. As the brain-cells multiply, the
prosencephalon outgrows the associated parts, and becomes nearly
separated into lateral halves ; these are the hemispheres of the
cerebrum, or “ halves of the great brain ” ; they retain their ven-
tricles, which intercommunicate through a passage-way, which also
leads into the third ventricle ; this is the foramen of Monro. Each
sends out in front a hollow process ; these processes are the olfactory
lobes, or rhinencephalon (“nose-brain”). A great ganglionic thicken-
ing of gray matter in the interior of each hemisphere is the corpus
striatum; these “striped bodies” are connected by the anterior com-
missure of the brain. The rest and greater part of the original
anterior cerebral vesicle makes up by ganglionic thickening of its
sides into what are called misleadingly the optic thalami, since these
tracts have nothing to do with the sense of sight. The thalami and
associate parts behind the lamina terminalis (third ventricle, etc.)
compose what some call the thalamencephalon, or “ bed-brain,” others
the diencephalon, or “ ’tween-brain.” The original middle cerebral
vesicle makes up underneath into longitudinal commissural fibres,
called the crura cerebri, or “legs of the brain,” connecting fore and
aft parts ; but especially composes the ganglionic centres called cor-
pora bigemina, or “ twin bodies.” These are the optic lobes, or “ eye-
brain.” They are connected by transverse commissure. The optic
ganglia and commissure, the cerebral crura, and contained cavities,
essentially compose the mesencephalon, or “mid-brain.” The original
posterior cerebral vesicle (opisthencephalon) becomes separated into
two parts : the fore part of it is moulded into the considerable mass
of the cerebellum (“ little brain ”) ; which, with its connections of

26o

GENERAL ORNITHOLOGY

PART II

white substance (pons Varolii, peduncles, etc.) and the hollow under-
neath it (“ fourth ventricle ”) constitutes the epencephalon of most
writers, by some called the metence;phalon^ or “after-brain.” The
hiud part of it tapers off into the spinal cord ; this tapering part
is the medulla oblongata, or “ oblong marrow,” also called the myelen-
cephalon, or “ marrow-brain ” (and by some the metencephalon). This
description is pertinent to brains at large, representing the general
plan of structure ; any fairly developed encephalon shows the parts
specified ; and the most complicated brain, that of man, only shows
what elaborate finishing touches may be given to the simple struc-
ture thus outlined, when cells, both white and gray, but especially
the latter, are profusely furnished, to the ornamentation of the
mind’s estate with race-tracks great and small, and the place of for-
nication,— fruits of the olive, and of the arbor vitae. The mem-
branes, or meninges, which hide all this from the uninitiated, are
three. The pia mater, or “tender mother,” which immediately
invests the brain, is very vascular, and furnishes the blood supply ;
not only by small arteries which immediately penetrate the substance
of the brain, but by enfolded sheets which enter the ventricles, and
are called choroid plexus. The arachnoid, or “ cobweb,” comes next ;
a serous fluid which it secretes bathes the brain, and meets con-
cussion with its gentler fluctuation. The dura mater, or “stern
mother,” is a dense outer membrane which enwraps and holds the
whole firmly. These meninges descend into the spinal column, and
answer the same purpose there, maintaining the same disposition
around the spinal cord.

The Bird’s Brain offers the following comparative characters : It
is compact, having nothing of the straggling apart of its elements
seen in low vertebrates, and completely fills the cranial cavity. Its
long axis is about transverse to the axis of the spinal column. The
cerebral hemispheres are well developed, but do not cover the cere-
bellum or optic lobes ; from their dome the rhinencephalon protrudes
like a porte-cochere. Their surface is quite smooth (devoid of the
gyri and sulci of most mammalian brains) ; even the Sylvian fissure
is barely indicated. The optic lobes are of immense size, relatively
to those of higher vertebrates, and relatively to the rest of the en-
cephalon ; they appear much loosened from their surroundings, at the
sides and lower part of the mid-brain ; they retain their ventricles, as
does also the rhinencephalon. The corpora striata are very large.
The fornix is rudimentary. The cerebellum is well developed and
deeply silicate, with transverse fissures, but is not divided into right
and left lobes ; a “ fleecy ” lobule on each side, the flocculus, is well
defined, and received in a special recess of the inner wall of the
skull. Parts of the medulla oblongata notable in mammals are
obscure or obsolete. There is no pons Varolii, or superficial trans-

SEC. IV

ANATOMY OF BIRDS

261

verse commissure of the cerebellum, nor any corpus callosum — that
great white commissure of the cerebral hemispheres which is
characteristic of all but the lowest mammals.

The Spinal Cord, or medulla spinalis (“spinal marrow”) is the
main nerve-axis of the body, running in the series of neural arches
of the vertebrae from head to tail ; it directly continues the medulla
oblongata. It retains its primitively tubular character in part at
least, and consists as usual of white matter enclosing gray matter.
The cord is fissured into lateral columns, as these are also to some
extent into anterior and posterior tracts. The latter diverge in
ascending the medulla oblongata, to throw the central tube into the
cavity of the fourth ventricle j and especially in the sacral region,
where a sort of ventricle, known as the avian sinus rhomboidalis, is
similarly formed. The calibre of the cord increases at the root of
the neck, where large nerves are to be given off from the brachial
plexus to the wings, and again in the sacral region, with the same
reference to nerve-supply of the legs ; after which the cord con-
tinues to the end of the spinal canal as a terminal thread.

The Cranial Nerves are twelve pairs, as in mammals, the
highest vertebrate number. 1, the olfactory nerve of special sense
(smell) ; origin from rhinencephalon ; exit from cranial cavity by
olfactory foramen, high up in orbital cavity ; conducted along a
groove to final escape between perpendicular and lateral plates of
ethmoid into the nasal chambers ; distributed to the investing
mucous membrane of the septal and turbinal bones of the nose.
The exit is through a sieve-like or cribriform plate only in Apteryx
and Dinornis (Owen). 2, the optic, nerve of special sense (sight) ;
origin from optic lobe and thalamus ; of great size, and forming a
chiasm (decussation) with its fellow ; exit by optic foramen, a large
hole in back of orbital cavity between centres of orbitosphenoid and
alisphenoid, close to or in common with its fellow. This nerve
forms the retina of the eye. 3, 4, 6, the oculimotor, pathetic, ab-
ducent, collectively the motor nerves of the eye, supplying the
muscles moving the eyeball; 3, to all these muscles excepting
superior oblique, and external rectus ; origin from crura cerebri, base
of mesencephalon ; 4, to the superior oblique, origin behind optic
lobes, upper surface of metencephalon ; 6, to external rectus (also
to muscles of the third eyelid in birds) ; origin between met- and
myel-encephalon, base of brain ; 3, 4, 6, exits from cranial into
orbital cavity by several small, not constant, foramina near optic
foramen ; or by this foramen sometimes all the nerves which enter
the orbit pass out of brain cavity through one great hole. 5, great
trifacial or trigeminal, sensorimotor ; feeling skin of head, moving
muscles of jaws ; origin (double) from myelencephalon ; leaves brain
from sides of metencephalon ; sensory root has Gasserian ganglion ;

262

GENERAL ORNITHOLOGY

PART II

motor root simple. This nerve has three divisions, whence its
name ; ophthalmic division, the most distinct ; exit from cranial
into orbital cavity by separate foramen above and to outer side of
optic foramen ; grooves orbital wall in passing ; ciliary ganglion ;
distribution mainly to lacrymal and nasal parts ; traceable to end of
upper mandible \ 5b, superior maxillary ; exit by foramen ovale, in
alisphenoid or between that and prootic centre ; distribution to side
of upper jaw ; Meckelian ganglion ; 5c, inferior maxillary, derived
chiefly from motor root ; exit same as 5h ; distribution to lower
jaw (muscles, substance of bone, integument) ; no special sense (gus-
tatory) function ; no otic ganglion. 7, facial or portio dura, motor ;
origin from myelencephalon ; enters periotic bone, escapes from ear
behind quadrate bone, by what corresponds to stylomastoid fora-
men of mammals ; communicates with 5c by chorda tympani nerve,
with 9, 10, 12, and sympathetic system; distribution to skin-
muscles and others of lower jaw and tongue, etc. 8, auditory or
portio mollis, nerve of special sense (hearing) ; origin with 7 ; no
exit from skull ; enters meatus auditorius internus of periotic bone ;
forms auditory apparatus in labyrinth of ear. 9, glossopharyngeal,
mixed nerve, sensorimotor and gustatory (taste) ; origin myelence-
phalon ; exit by foramen in exoccipital bone, behind basitemporal,
near lower border of tympanic recess ; distribution to muscles and
membranes of gullet, throat, tongue, etc. \0, pneumogastric, sensori-
motor ; origin and exit next to 9 ; distribution to windpipe, lungs,
gullet, stomach, heart, etc. ; has recurrent syringeal to vocal organs.
11, spinal accessory, sensorimotor; origin upper part of spinal cord ;
exit with 9, 1 0 ; distribution to these nerves and to muscles of
neck. 9, 10, 11, are intimately connected with one another, and
with other nerves, especially 10 with sympathetic. The several
foramina in a bird’s skull, which may be seen in the place indicated
at 8, Figs. 69, 70, are for the divisions of this composite vagus or
“ wandering ” nerve of respiration, circulation, digestion, etc. ; they
represent morphologically a foramen lacerum posterius, between ex-
occipital and opisthotic centres. 12, hypoglossal, motor nerve of the
tongue ; origin from myelencephalon ; exit by anterior condyloid
foramen in front of the occipital condyle. Thus the plan of the
cranial nerves of birds is nearly coincident with that of mammals.

The Spinal Nerves, in pairs, correspond in a general way to
the vertebrae, between which they pass out by intervertebral fora-
mina, to supply the body at large. They are sensorimotor ; arise
from the spinal cord by anterior motor and posterior sensory
(ganglionated) roots which unite before leaving the spinal canal ; in
the sacral region the main branches leave by separate foramina.
They form plexuses or interlacements. The principal of these is the
brachial plexus ; constituted by several lower cervical nerves, and

SEC. IV

ANATOMY OF BIRDS

263

one or two usually counted as dorsal, which combine to form a
single cord, whence the nerves of the wing are derived. Similar
network of three to five true sacral nerves furnishes the nerves of
the leg.

The Sympathetic System consists of a pair of nervous cords
running lengthwise below the bodies of the vertebrae, one on each
side in the trunk, and in corresponding relations with cranial bones.
An extensive and intricate series of communications is effected with
the nerves of the cerebrospinal system, excepting the special-sense
nerves of smell, sight, and hearing. The points of communication
form a chain of sympathetic ganglia ; from these knots, the most
conspicuous features of the system, nervous cords pass to their dis-
tribution in the motory mechanism of the heart and blood-vessels
and other viscera. The anterior sympathetic nerves are the iridian;
the anterior ganglia are the SThenoTalatine or Meckelian^ intimately
connected with cranial nerves. The system ends behind in the
caudal region of the spine by a ganglion impar.

Sense of Smell : Olfaction. — The sense of smell is effected by
terminal branches of the olfactory (1st cranial) nerve, ramifying in
the mucous (pituitary or Schneiderian) membrane of the nasal
cavities. Owing to the comparatively small size and little com-
plexity of the foldings and pleatings of bone or cartilage in the
nasal chambers, the sensory surface being correspondingly limited,
it is not probable that birds possess this sense in a high degree.
Besides the cartilaginous or osseous septum, generally more or less
complete in birds, there are lateral scrolls and whorls of bone in
endless diversity in most birds, which may be ossified, or remain
gristly. The general cavity is mostly bounded and enclosed by the
bony beak ; floored by the anterior part of the hard palate ;
defended on each side by the descending prong of the nasal bone ;
in the dry skull, it either seems continuous with the great orbital
cavity on each side behind, or is separated therefrom by lateral eth-
moid (prefrontal) or lacrymal ossifications, or both. Outwardly the
nasal chambers open upon the beak by the external nostrils. These
openings are minute or quite obliterated in some Steganopodes, as
pelicans and cormorants. The nasal cavities always communicate
with the back part of the mouth by the posterior nares (Lat. naris, a
nostril) ; they are generally paired, that is, with a partition between
them, sometimes united in one median aperture. The olfactory
nerve, which is rather a prolongation of the rhinencephalon itself
than an ordinary nerve, escaping from the brain-box by a special
foramen, traversing the upper part of the interorbital septum in a
groove or canal, enters the nasal cavity by a single orifice (excepting
Apteryx and Dinornis), instead of the numerous apertures in a cribri-
form plate by which its filaments reach their destination in mam-

264

GENERAL ORNITHOLOGY

PART II

mals. The true sensitive membrane, in which the nervous filaments
end, is that investing ethmoidal (septal and turbinal), not maxillary
parts. An associate structure of the olfactory organ is the nasal
gland, sometimes called the sujgerorhital gland, from its position in
many birds. Thus it is of great size in a loon, and lodged in a
large deep crescentic depression on top of the skull over the orbits
(Fig. Q3,w) ; these crescents nearly meeting each other in the middle
line. In other birds it is smaller, and within the cavity of the
orbit, but never in that of the nose itself, its secretion being
poured into the nasal chamber by a special duct.

Sense of Sight : Vision. — The eye is an exquisitely perfect
optical instrument, like an automatic camera obscura which adjusts
its own focus, photographs a picture upon its sensitised retinal
plate, and telegraphs the molecular movements of the nervous sheet
to the optic “ twins ” of the brain, where the result is “ biogenised ” ;
that is, translated from the physical terms of motion in matter to
the mental terms of consciousness. But no part of the nervous
tract, from the surface of the retina to the optic centre, sees or
knows anything about it, being simply the apparatus through which
the Bird looks, sees, and knows. In this class of Vertebrates, the
optic organs, both cerebral and ocular, are of great size, power, and
effect ; their vision far transcends that of man, unaided by artificial
instruments, in scope and delicacy. The faculty of accommodation,
that is, of adjusting the focus of vision, is developed to a marvellous
degree : rapid, almost instantaneous, changes of the visual angle
being required for distinct perception of objects that must rush into
the focal field with the velocity at least of the bird’s flight. Birds
are therefore far-sighted or near-sighted (presbyopic or myopic)
according to the degree of tension the nerve-tide excites in the eye
by the mechanism described farther on ; and the transition from
one to the other state is effected with great quickness and correct-
ness. Observe an eagle soaring aloft until he seems to us but a
speck in the blue expanse. He is far-sighted ; and scanning the
earth below, descries an object much smaller than himself, which
would be invisible to us at that distance. He prepares to pounce
upon his quarry ; in the moment required for the deadly plunge he
becomes near-sighted, seizes his victim with unerring aim, and sees
well how to complete the bloody work begun. A humming-bird
darts so quickly that our eyes cannot follow him, yet instantaneously
settles as light as a feather upon a tiny twig. How far off it was
when first perceived we do not know ; but in the intervening frac-
tion of a second the twig has rushed into the focus of distinct
vision, from many yards away. A woodcock tears through the
thickest cover as if it were clear space, avoiding every obstacle.
The only things to the accurate perception of which birds’ eyes

SEC. IV

ANATOMY OF BIRDS

265

appear not to have accommodated themselves are telegraph-wires
and lighthouses ; thousands of birds are annually hurled against
these objects to their destruction.

The orbital cavity, orbit, or socket of the eye, has been almost
sufficiently described (see also any Figs, of skull in profile) as that
great recess in the side of the skull which is bounded above by the
roofing frontal bone, behind by this and sphenoidal elements, in
front, if at all, by lateral ethmoidal elements (prefrontal), and
separated from its fellow more or less completely by the inter-
orbital septum, which is chiefly the perpendicular plate of the
mesethmoid, but may be also in part orbitosphenoidal and pre-
sphenoidal. The brim is completed in few birds, by union of lacry-
mal and postfrontal ; in quite a number of birds, however, it is
nearly perfected by the approximation of these same bones, as in
Fig. and m, and in some the rim is carried out by extra supra-

orbital and infraorbital ossification. There is no bony floor, or
only such slight scaffolding as the expansion of the palatine and
pterygoid may afford. The zygoma itself, in many dry skulls,
seems like the threshold of the orbital chamber. The bony walls
may be also defective in some places by great vacuities in the inter-
orbital septum (Fig. 70, iof, and Fig. 63, z), and others in the cerebral
wall, aside from the regular foramina which the nerves pass through.
The 1st — 6th nerves (p. 261) inclusive usually enter the orbit : of
their foramina, the (Figs. 66, 68, 70, 71, and Fig. 63, y)
is much the largest and most constant, generally blended with its
fellow. Those for nerves 1 and 5 (p. 261 ) are next most obvious and
constant ; others are often, and all may be, thrown into one large
opening. In such a socket as this the eyeball rests upon a cushion
of muscle, fat, gland, and connective tissue ; and large as is the
chamber, the ball fits and nearly fills it. A bird’s eyeball is much
larger than the opening of the eyelids (see p. 45, note 3).

As to its development : “ the Eye,’^ says Huxley, “ is formed by
the coalescence of two sets of structures, one furnished by an involu-
tion of the integument, the other by an outgrowth of the brain.
The opening of the tegumentary depression, which is primarily [in
the very early embryo] formed on each side of the head in the
ocular region, becomes closed, and a shut sac is the result. The
outer wall of this sac becomes the transparent ccrrnca of the eye ;
the epidermis of its floor thickens, and is metamorphosed into the
crystalline lens ; the cavity fills with the aqueous humour. A vascular
and muscular ingrowth taking place round the circumference of the
sac, and dividing its cavity into two segments, gives rise to the iris.
The integument around the cornea, growing out into a fold above
and below, results in the formation of the eyelids, and the segrega-
tion of the integument which they enclose, as the soft and vascular

266

GENERAL ORNITHOLOGY

PART II

conjunctiva. The pouch of the conjunctiva very generally communi-
cates, by the lacrymal duct, with the cavity of the nose. It may be
raised, on its inner side, into a broad fold, the nictitating membrane,^
moved by a proper muscle or muscles. Special glands — the lacrymal
externally, and the Harderian on the inner side of the eyeball — may
be developed in connection with, and pour their secretion on to, the
conjunctival mucous membrane. The posterior chamber of the eye
has a totally distinct origin. Very early that part of the anterior
cerebral vesicle, which eventually becomes the vesicle of the third
ventricle, throws out a diverticulum, broad at its outer, narrow at
its inner end, which applies itself to the base of the tegumentary
sac. The posterior, or outer, wall of the diverticulum then becomes,
as it were, thrust in, and forced towards the opposite wall by an
ingrowth of the adjacent connective tissue ; so that the primitive
cavity of the diverticulum, which, of course, communicates freely
with that of the anterior cerebral vesicle, is obliterated. The broad
end of the diverticulum acquiring a spheroidal shape, while its
pedicle narrows and elongates, the latter becomes the optic nerve,
while the former, surrounding itself with a strong fibrous sclerotic
coat, remains as the posterior chamber of the eye. The double
envelope, resulting from the folding of the wall of the cerebral optic
vesicle upon itself, gives rise to the retina and the choroid coat, the
plug or ingrowth of connective tissue gelatinises and passes into the
vitreous humour, th.Q cleft by which it entered becoming obliterated.’’
{Anat. Vert, 1871, p. 79.)

Birds alone, of all animate beings, may be truly said to “ fall
asleep ” in death. When the “ silver cord ” of a bird’s life is loosed,
the “ windows of the soul ” are gently closed by unseen hands, that
the mysterious rites of divorce of spirit from matter may not be
profaned. When man or any mammal expires, the eyes remain
wide open and their stony stare is the sign of dissolution. Only
birds close their eyes in dying. At the same moment, the eye
sinks and seems to collapse, by the ebbing of its waters. The
closure is chiefly effected by the uprising of the lower lid. These
are the principal external differences between the eyes of birds and
mammals. The movements of the upper lid in most birds are much
more restricted than those of the lower. The few exceptions are
chiefly furnished by night birds, as owls, whip-poor-wills, and others
of their respective tribes. The lids consist externally of common
skin, internally of a layer of conjunctival (joining) mucous membrane,
with interposed connective tissue : the lower is also stiffened with
a smooth plate, the tarsal cartilage. The upper is raised by a small
muscle, called from its office levator palpehrce superioris, arising from
the bony orbit. There is no special lowering nor lifting muscle of
the under lid ; the lids close together by the action of the orbicularis

SEC. IV

ANA TOMY OF BIRDS

267

ocul% which nearly surrounds the eye, and whose chief office is to
lift the lower lid ; the latter has a small distinct depressor muscle.
Birds have no true hairs, but in some kinds modified filiform feathers
answer to eyelashes. When wide open the orifice of the lids is
circular, that is, without the inner and outer corners (canthi) of
almond-eyed creatures like man. There is a third inner eyelid,
highly developed and of beautiful mechanism : this is the nictitating
membrane, or “winker” (nictito, I wink), a delicate, elastic, trans-
lucent, pearly-white fold of the conjunctiva. While the other lids
move vertically and
have a horizontal com-
missure, the winker
sweeps horizontally or
obliquely across the ball,
from the side next the
beak to the opposite.

If we menace a bird’s
eye with the finger, it
is curious to see the
winker rush out of the
corner to protect the
ball. Owls habitually sit
in the daytime with this
curtain shading the eyes
from the glare of light ;
and doubtless the eagle
throws the same screen
over its sight when soar-
ing towards the sun.

•x-rj-l i • . • .1 lUUSUlCS . ttj OUjJCllUl , Uf i.CU;ijU.O CA.OClilU.0 , u,

W nen not in action, tlie rectus inferior ; d, rectus internus ; e, obliquus superior ;
winlrpr lipct pnrlpd nn in lettered), obliquus inferior; 9?, quadratus ; h, pyra-

WinKer lies curietl up in n^i^alis, with its tendon, fc, passing through a pulley in
the corner of the eve the quadratus (as shown by the dotted line) to keep it off
, ’ the optic nerve, i, then passing around the edge of the

like those patent window bail to its insertion in the nictitating membrane.

shades which stay up of

themselves till pulled down. The ingenious mechanism of the
movement of the winker across the lid may be understood with the
help of Fig. 81, which represents the hao^k of the eyeball. The
winker lies in front, on the left hand of the picture, and is to be
pulled across the front by the slender tendon, k, of the pyramidalis
muscle, h. As h contracts it pulls on k, and winding round to
the front, pulls the winker to the right hand. But i is the
optic nerve, entering the ball ; k would press upon it, were it
not fended off by passing, as seen by the dotted line, through a
pulley in the end of the quadratus muscle, g. The harder h pulls,
the harder does g also pull, their consentaneous action at once

Fig. 81. — Right eyeball, seen from behind, showing
the muscles : a, rectus superior ; 6, rectus externus ; (

268

GENERAL ORNITHOLOGY

PART II

giving the proper direction to the tendon and keeping it off the
nerve.

Beneath the eyelids, upon the ball, is a delicate filmy membrane
not easily recognised on ordinary inspection : this is the conjunctiva^
so called because it joins the eye to the lids. The ocular layer is
transparent where it passes over the cornea : it is then reflected
away from the ball, to form the ]palpebral layer, — a folding between
being the nictitating membrane. The conjunctiva is highly vascular,
but the blood-vessels are too small to be seen unless they become
congested, when the eye presents the well-known appearance called
“ blood-shot.” Though birds can hardly be said to cry, they have a
well-developed apparatus for the manufacture of tears. The lacrymal
are two small glands lying one in each corner of the eye, inner and
outer. The former, called the Harderian gland, is the smaller, deeply
seated behind the winker, upon which it pours a glairy fluid : it is
an oil-can which not only supplies but applies the fluid to the
winker, w^hich needs constant lubricating to work well. The lac-
rymal gland proper is the outer one, which prepares the tears to
moisten and cleanse the conjunctiva ; after which they are drained
off by the lacrymal duct into the cavity of the nose, which thus
becomes a sort of cesspool to receive the refuse waters of the eye.
A third gland about the orbit has been already mentioned (p. 264)
as pertaining to the nose, not to the eye. Its site is shown in the
crescentic superorbital depression. Fig. 63, w.

The motions of the eyeball, though more restricted than in
mammals, owing to the shape of the ball and its close socketing,
are nevertheless subserved by the usual number of six muscles. Of
these four are called the recti, or straight muscles, and two the
ohliqui, or oblique muscles ; though they are all “ straight ” enough,
the terms applying to their lines of traction. The four recti arise
from the bony orbit, near together, about the optic foramen, and
pass to be inserted in the eyeball at as many nearly equidistant
points on its circumference; the musculus rectus superior, Fig. 81,
on top ; m. r. inferior, c, below, antagonising a ; the m. r. externus,
b, and internus, d, respectively to the outer and inner (hindward
and forward) sides, also antagonising each other. The two oblique
muscles arise farther forward in the bony orbit, near each other,
and then diverge obliquely upward, m. o. superior, e, and downward,
m. 0. inferior, f, to be inserted near the margin of the globe of the
eye, close by the respective insertions of superior and inferior rectus.
All the motions of the ball result from consentaneous or dissen-
taneous action along these six lines of traction ; the muscles acting
as ropes to pull the ball about, and to steady it in any direction
of its axis. The peculiarity of mechanism in a bird is, that the
superior oblique goes straight to its insertion, instead of passing

SEC. IV

ANATOMY OF BIRDS

269

through a pulley which changes its line of traction in mammals.
The special nerves presiding over these muscles (3, 4, 6) have been
pointed out already (p. 261). In the figure, the cut orbital ends of
them all are reflected away from the ball to disclose the underlying
muscles of the winker: the reader must mentally bring the six
loose ends together and fasten them to the bony orbit at points
near about opposite i, as above said of their origins.

The above are the principal circumstances and accessories of the
optic apparatus ; we may now examine the eye itself, of which Fig.
82 gives an enlarged view, in longitudinal vertical section, — the

Fig. 82. — Vertical antero-posterior section of eyeball ; a, optic nerve ; 6, sclerotic, its outer
coat ; c, sclerotic, its middle and inner coats ; d, choroid ; e, hyaloid ; /, marsupium ; g, cornea ;
h, Ji, bony plates between sclerotic layers ; i, i, corrugations of choroid, forming ciliary pro-
cesses ; k, k, canal of Petit ; I, I, iris ; m, anterior chamber of eye ; n, capsule of the lens ; o,
lens ; p, posterior chamber of eye. Neither the retina, nor the peculiar sheathing of the optic
nerve, is shown. The nerve, marsupium, and ciliary processes, not falling in this section, can
only be arbitrarily shown.

nerve, marsupium, and ciliary processes not indeed lying as shown
in this section, but so introduced as to display them intelligibly.
A bird’s eyeball is not nearly so spherical or globular as a mam-
mal’s. The globe of the human eye is about a five-sixths segment
of a large sphere (sclerotic) with a one-sixth segment of a smaller
sphere protruding in front (corneal). The anterior part of the
sclerotic of a bird is so prolonged as to be in some cases almost
tubular or cylindric, and the corneal protuberance is very convex :
the figure may be likened to an acorn which has a short blunt
kernel in a heavy shallow cup, or to a thick old-fashioned watch
with a very convex crystal. This characteristic shape is fairly shown

270

GENERAL ORNITHOLOGY

PART II

in the figure ; but some birds’ eyes are much more tubular in front,
— owls’, for example. The eyeball being hollow and filled with
fluids which press in all directions, it is hard to see at first how
such a peculiar shape is maintained. But the sclerotic coat is very
dense, almost gristly in some cases ; and it is reinforced by a circlet
of the sclerotals, A, h; see also Fig. 62, where the circlet is

shown. These are packed alongside each other all around the cir-
cumference of one part of the sclerotic, like a set of splints. The
large discoidal segment of a bird’s eye is mostly composed of the
membrane called from its hardness the sclerotic, — thick, tough, and
strong, of a glistening livid colour. Three sclerotic coats or layers
may be demonstrated by careful dissection ; in the figure h is the
outer, c the combined middle and inner ones, — much exaggerated
as to their distinctness. The bony plates lie between the outer and
middle coats anterior to the greatest girth of the eyeball, extending
from the rim of the disk nearly or quite to the edge of the cornea.
They are a dozen to twenty in number, of oblong squarish shape,
tapering toward the cornea, around which they are thus circularly
disposed ; they are pretty closely bound together, but the circlet as
a whole enjoys some little motion back and forward with the vary-
ing convexity of the cornea, g. This last is the thin transparent
membrane completing the eyeball in front, like the crystal over
the face of a watch. It is very protuberant in birds, — even a
hemisphere, or almost tubular. Its structure is not peculiar in
birds ; but it is remarkable in this class of creatures not only for
its convexity, but for the wide range of the variability in convexity
which increased or diminished pressure of the contained humours
may effect, and its collapse in death.

The sclerotic coat is lined with the choroid membrane, d, loosely
woven of cellular tissue, replete with blood-vessels, and painted
pitch-black with a heavy deposit of pigment-cells. It lines the
whole globe as far forward as the edge of the sclerotal bones, where
it splits in two layers. The inner choroid layer turns away from
the wall of the eye, toward the interior, and in so reflecting becomes
plaited, as a bag is puckered by pulling the strings. These pleats
converge upon the rim of the delicate capsule enclosing the lens of
the eye, n, and there adhere, forming the ciliary processes, i, i The
outer layer also starts away from the circumference of the sclerotic
wall, as if to pass directly across the cavity, but ends in the iris.
Around the circumference of the iris, where sclerotic, corneal, and
choroid coats come together, is a circular band of fibres, the ciliary
ligament ; and on the outer surface of the choroid is a similar band
of circular and radiating contractile fibres, the ciliary muscle. These
ciliary structures are supposed to be the agents of the accommodat-
ing faculty of the eye, acting upon the lens to alter its shape or its

SEC. IV

ANATOMY OF BIRDS

271

position, or both. This is a difficult matter to settle, when such
delicate structures are in question.

The m5, /, Z, or rainbow of the eye, is an exquisite structure
hanging like a many-coloured curtain vertically between the two
compartments of the eye ; a highly ornamental framework of the
eye’s window, being both sash and blind to the pupil. It is sus-
pended vertically in the aqueous humour, just in front of the lens.
Viewed in front, from the outside, the iris appears as a coloured
circular band around the pupil, and seems to come to the surface of
the eye. But this is not so, for the conjunctiva, the cornea, and
the aqueous humour of the front chamber of the eye, are between
us and it. It may be likened to the dial-plate of a watch, which we
look at without noticing the interposed crystal. Similarly, the pupil
of the eye, which shows us our own reflection, diminished to the
size of the “ eye-baby,” may be likened to the round central hole in
the dial-plate through which protrudes the shaft that bears the
hands of a watch. The “ pupil ” is the round black spot within
the coloured rim of the iris ; but it is not a thing — it is a hole in a
thing — the hole in the iris through which we may look and see the
black choroid coat behind. The quivering iris is very similar in
texture to the choroid, being a delicate tissue of interlacing fibres
and vessels ; but it is highly mobilised by circular and radiating sets
of contractile fibres, by which the curtain is tightened and loosened,
with corresponding change in the size of the central orifice — the
pupil. Although the iridian movements are largely automatic,
depending upon the stimulus of light, they are to some extent
voluntary, as any one may satisfy himself who observes owls in con-
finement. During these expansions and contractions of the iris the
pupil in birds preserves its circularity ; and even when the move-
ment is freest and most voluntary, as in owls, the contracted pupil
never appears as a vertical oval figure, or a slit, like that of cats.
The round pupil of the great horned owl. Bubo virginianus^ ranges
from the diameter of a finger-ring down to that of a small split-
pea. The iridian colours are often striking in birds. Though
black and brown are the commonest, yellow is quite frequent, red
is often seen, blue and green are rarer; the eyes of cormorants
are of the latter colour. The iris is sometimes pure white, as it is
in the white-eyed greenlet, Vireo noveboracensis. In the Californian
woodpecker, Melanerpes formicivorus, the eyes are indifferently (or
at different ages of the bird, or seasons) brown, bluish, pink, rosy,
or yellow.

The crystalline lens, 0, is a transparent biconvex disk like a com-
mon magnifying glass, apparently set in the iris like a mirror in its
frame, but really hanging a little back of that structure. It is
enclosed in a capsular membrane, n, of extreme delicacy and trans-

272

GENERAL ORNITHOLOGY

PART II

parency, which is in turn set between two layers of the hyaloid
membrane to be presently noticed. Where these layers of hyaloid
separate around the rim of the capsule to form the investment, a
small space is left between them ; this circular tube around the lens
is the canal of Petit, k, k. The lens is stationed in the axis of
vision ; some suppose it to be equally stationary in any transverse
axis. It is, however, difficult to understand how an object thus
suspended in fluctuating humours should be insusceptible of some
motion backward or forward as well as of alteration in its degree of
convexity ; both of which may be factors in the focusing process.
From what has preceded, it is evident that the cavity of the eye is
divided into anterior and posterior compartments, or chambers, by
the reflection, from the sclerotic wall, of the choroid, hyaloid, and
iridian structures, which with the lens form a vertical partition.
Each chamber is filled with a fluid of different density and con-
sistence. That in the anterior or corneal chamber is thin and
watery, and therefore called the aqueous humour; that in the sclerotic
cavity is more dense and glassy, and for this reason known as the
vitreous humour. There is much less aqueous than vitreous ; but
birds have comparatively more of the former than usual, owing to
the relatively greater size and convexity of the cornea. The waters
are enclosed in exceedingly delicate membranes ; the vitreous in the
hyaloid membrane, e, which, besides lining the posterior chamber and
enclosing the lens as already said, sends thin partitions all through
the vitreous humour to steady these glassy waters.

The optic nerve, a, of birds is peculiar. In mammals, as a rule,
the nerve is a smooth cylinder, proceeding straight to the sclerotic,
penetrating the coats of the eyeball directly, near the middle point
behind, and then spreading out on the inside of the ball as a large
circular concave mirror. This thin, saucer-like expansion of nerve-
tissue is the retina. In birds the optic nerve is a fluted column,
which approaches the eyeball quite obliquely, strikes it at a point
eccentric from the axis of the eye, and does not at once pierce the
sclerotic. Tapering to a fine point, and running still obliquely,
downward and forward, in a deep groove in the sclerotic that would
be a tube were it not split, and through a similar slit in the choroid,
a fluting of the nerve rises to attain the cavity of the eye, and the
retina spreads out from the sides and end of this fold. But the
prime peculiarity of a bird’s eye is the “ purse ” or “ comb,” mar-
supium, pecten, //a very vascular structure, like the choroid, and
likewise painted black ; apparently “ erectile,” that is, capable of
increasing and diminishing in size by influx and efflux of blood. It
is attached behind to the nervous structure ; is suspended in the
vitreous humour, and runs forward obliquely a part or the whole of
the way to the lens, to the envelope of which it may be attached in

SEC. IV

ANATOMY OF BIRDS

273

some cases. Its office is not fully determined. Its great resem-
blance to the choroid proper suggests a similar function in the
absorption of light. If it be turgid and flaccid by turns it must
occupy a variable space in the vitreous humour, and in the former
state press the waters upon the most yielding part of their walls, —
that where the lens is situated, even to the extent of altering the
position of the latter ; and if so, of changing the focus of the eye. It
is difficult to account for the bird’s eyes’ powers of accommodation
by the action of the ciliary muscle in only changing the sha'pe, of the
lens, thus throwing out of account as impossible any change in the
position of that refracting medium, or of the density of the refracting
humours, or of the convexity of the cornea. The peculiar course
of the optic nerve may be simply an anatomical convenience, or may
have something to do with a bird’s ability to see straight ahead
though its eyes be laterally positioned. (See Am. Nat. ii. 1868, p.
578; Pr. Post. Soc. Nat. Hist. xii. Apr. 21, 1869.)

Sense of Hearing : Audition. — This is enjoyed to a high degree
by the “musical class” of the Vertehrata, — birds being the only
animals besides man whose emotions are habitually aroused,
stimulated, and to some extent controlled, by the appreciation of
harmonic vibrations of the atmosphere. Most birds express their
sexual passions in song, sometimes of the most ravishing quality to
our ears, as that of the nightingale or the bluebird, and it cannot
be supposed that they themselves do not experience the effect of
music in an eminent degree of pleasurable perturbations. Other-
wise they would cease to sing. The capability of musical expression
resides chiefly in the more spiritualised male sex ; the receptive
capacity of musical affections is better developed in the female, who
chiefly furnishes the plastic material which is to be moulded into
the physical manifestation of the male principle. Quickness of ear
is extraordinary in such birds as those of the genus Mimus, which
correctly render any notes they may chance to hear with greater
readiness and accuracy than is usually within human possibility. It
may be reasonably doubted that any others than some of the world’s
greatest musical composers have a higher experience in acoustic
possibilities than many birds. Birds’ ears have nevertheless a com-
paratively simple anatomical structure, on the whole much more like
that of reptiles than of mammals. Such simplicity is seen in the
ligulate or strap-shaped cochlea, the essential organ of hearing. Figs.
84, 85, 86, 87, as compared with the helicoid curvation of the
mammalian cochlea. The openness of the ear -parts which lie
outside the tympanum is seen in Fig. 62, at the place where the
reference-lines “ ear-cells ” reach the skull ; and especially in Fig. 71,
where the stapes, st, is seen lying in the ear-cavity, the tympanum
having been removed.

T

274

GENERAL ORNITHOLOGY

PART II

There is ordinarily no external ear, in the sense of a fleshy
conch or auricle, though owls at least have a considerable flap which
overlies the auditory aperture. The place of an auricle is filled by
a set of peculiarly modified feathers surrounding and overlying the
ear -opening, called in ornithology the ear -coverts, or auricular s
(Fig. 25, The outer ear or meatus auditorius externus is a con-
siderable shallow roundish depression in the skull, at the extreme
lower lateral corner. Its ordinary boundaries are the movably
articulated quadrate bone in front, the expanded rim of the squa-
mosal above, the tympanic wing of the exoccipital behind and below ;
the termination of the basitemporal also usually contributing to the
under boundary. (See Fig. 71, at sf ; 63, under I ; Fig. 62, where
reference lines “bones of ear-cell” go.) On removing the quadrate
from the dry skull the general tympanic depression is seen to be
more or less continuous with the alisphenoid ; the boundary is best
marked behind and below by the broad thin sharp-edged shell of
the tympanic wing of the exoccipital. To the brim indicated is
attached the tympanum, or drum of the ear — that membrane being,
from the configuration of the parts, quite superficial, — not at the
bottom of a tube-like meatus, as in man. The membrane proper is
invested externally by modified common integument which readily
peels off. Thus this wide shallow depression overlaid with feathers
or a slight flap is all there is to represent the “ outer ear-passage.”
The tympanic membrane sometimes develops slight ossification,
which then represents the “ tympanic bone,” or “ external auditory
process ” of human anatomy. Did not this membrane occlude the
way, the passage through the ear to the mouth would be pervious.
This passage is the modified persistence of the first visceral cleft or
“ gill-slit ” of the embryo. Just within the tympanic membrane is
the cavity of the tympanum or middle ear, which may be very exten-
sively exposed by merely removing the membrane. Looking into
this cavity, as may readily be done from the outside, in carefully
cleaned dry skulls, many objects of interest are presented ; among
them, a number of foramina — openings leading in various directions.
In the first place there are some (inconstant and not readily iden-
tified) holes, which ate pneumatic openings, conveying air from the
middle ear-passage to the interior of bones of the skull and lower
jaw. Next is observed a large orifice in the lower anterior part of
the cavity, — the mouth of the Eustachian tube. This tube continues
the ear-passage to the mouth, and opens at the back of the hard
palate by a median orifice in common with its fellow. In clean
skulls of any size a bristle, or even a wooden toothpick, will pass
through the Eustachian tube, and appear upon the floor of the skull
in mid-line or nearly there, under the basisphenoid, over the basi-
temporal. The foregoing passages have not conducted us to the

SEC. IV

ANATOMY OF BIRDS

275

inner ear or proper acoustic cavity.. There will he observed, in the
side-wall of the tympanic cavity, two definite openings near the
Eustachian orifice. One of these, anterior and superior to the other,
larger usually, and oval, is the fenestra ovalis ; it lies in the ob-
literated suture between the prootic and opisthotic bones ; and when
the membranous curtain which closes it in life is gone, you look
through this “ oval window ” into the vestibular cavity of the ear
proper. The lower, posterior, circular orifice is the fenestra rotunda ;
through which round window in the opisthotic hone you look into
the cochlear cavity oi the ear proper. Fenestra ovalis and f. rotunda
are generally close together, — only divided by a little bridge of
hone, or a mere bony bar. To the circumference
of the fenestra ovalis is fitted the expanded oval
foot of the trumpet-shaped columella auris, — the
stapes, or “ stirrup-bone,” as it is called in mammals
(Fig. 83, st). This is an elegant little bone, which
establishes mechanical connection between the
membrane closing the fenestra ovalis and the
tympanic membrane, — something on the principle
of the “sounding-post” inside a violin. It is
shown magnified greatly in its embryonic condition
in Fig. 67, and there seems to be primitively and
morphologically the proximal connection of the
hyoid bone (by ceratohyal elements) with the bony
capsule of the ear ; but no trace of this relation
persists. Fisr. 83 shows the mature stapes of a ovalis; TOs^ main shaft,

f ® 1 . 1 1 01’ mediostapedial ele-

lowl, and indicates its several elements which have nient; sst, suprastape-

received special names. In skulls prepared with dial’; 7si, InflStapt'
sufficient care, the stapes may be seen in situ, as rudhneSry

in Fig. 71, st, — an extremely delicate rod, stepped styiohyai ;/, a fenestra
into the fenestra ovalis by its foot, the other end (see in^Juf^gl^nj
protruding freely, and bearing in many cases its mation,^Fig.’^67l^^^
hammer-like or claw-like stapedial elements. A
stapes I have just picked out of an eagle’s ear is a fourth
of an inch long, with a stout foot, but a stem as fine as a
thread of sewing silk, and at the tympanic end a still finer
hair -like process, half as long as the main stem, from which
it stands out at a right angle. The ossification is perfect, and there
appears to have been another similar process which has broken off
from the cross-like figure shown in Fig. 71, s^. In a raven’s skull
before me the stapes has fallen into the fenestra ovalis, and lies
there with its head sticking out. Though perfectly loose, I cannot
withdraw it intact, as the expanded foot fits the hole too closely to
pass through in any position I have succeeded in placing it. It
appears to be about as large as the eagle’s. Close examination at a

Fig. S3. — Mature
stapes of fowl, about x
4 ; after Parker, st, its
foot, fitting fenestra

276

GENERAL ORNITHOLOGY

PART II

point somewhere about the fenestra ovalis, or between that and the
Eustachian orifice, will discover a minute foramen corresponding
to the stylomastoid foramen of mammals. It transmits cranial
nerve 7 (see p. 262), or the facial nerve, which has burrowed through
the bony acoustic capsule from the brain-cavity and entered the
tympanic cavity on its way to the surface. There are sometimes
two such minute foramina, close together, both conducting to the
brain cavity (neither in common with the internal auditory meatus) ;
as in the eagle, in which large bird a fine bristle just passes through
each. Thus in the dry skull of a bird all the hard parts of the
middle ear or tympanic cavity, as well as the Eustachian tube, can
readily be inspected from the outside; even the limits of the
opisthotic and prootic bones can be determined to some extent, and
the ossiculum auditus be seen in situ. There will also be noted, in
most birds, the articular facet upon the prootic bone for the inner
head of the quadrate, as well as upon the squamosal for the outer
head of the quadrate; however these may shift in position, in
different birds, they cannot easily be overlooked or mistaken.
Details of mere size and configuration aside, the above general
description will apply pretty well to any bird, and should suffice
for the identification of the objects seen on looking into the ear,
though the number and variety of the irregular pneumatic openings
may be puzzling at first. To see these things clearly in a mammaVs
ear would require special preparation of the parts, as they lie inside
a tympanum which is itself at the bottom of a contracted tube. In
such an ear, properly laid open, would be found a chain of three
ossicles crossing the tympanic cavity from the inner surface of the
tympanic membrane to the opposite surface of the membrane closing
the fenestra ovalis — the malleus, incus, and stapes, or “hammer,”
“anvil,” and “stirrup”; and the latter would be stirrup-shaped,
not trumpet-like with a cross-bar at the mouthpiece. Some mam-
mals would also show a hyoid bone which would have what are
the ceratohyals of a bird produced up toward the ear-parts, and
continued to these by a bone called stylohyal, or “ styloid process of
the temporal ” ; and any mammal’s jaw would articulate directly
with the squamosal, — the chain of three ossicles being entirely inside
the ear. As to comparing the parts now : the mammalian stapes is
the stapes or columella of a bird, — its stem and foot at least ; the
incus of a mammal is represented by one of the claws of the cross-
bar of a bird’s stapes (the SMj?rastapedial element; Fig. 83, sst) ; the
malleus of a mammal is the great quadrate bone of a bird ; the
stylohyal of a mammal is not fairly developed in a bird, unless con-
tained in or represented by another claw of the stapes (an infra-
stapedial element, ist) ; and in these facts is the reason why a bird’s
lower jaw is articulated indirectly to the skull by means of the

SEC. IV

ANATOMY OF BIRDS

277

quadrate, and also why a bird’s hyoid bone is not articulated or in
any way directly connected with the skull — excepting when, as in
a woodpecker, elongated branchial elements of the hyoid bone take
on such office by curling over the cranium (Figs. 73, 74).

Section of the bone is required for further examination of the
ear-parts. On longitudinally bisecting the skull, or otherwise gain-
ing access to the brain-cavity, the internal surface of the ^periotic bone
is brought into view (Fig. 70, ^0, op, ep). It is the same bone we
have seen in the tympanic cavity, now viewed upon its cerebral
surface. In a skull of any size, as that of the eagle before me
(from which the rest of my description will be taken), there is no
difficulty in making out the parts, although the periphery of the
periotic bone is completely consolidated with its surroundings.
The periotic or petrosal (Lat. petrosus, stony — from its hardness), or
“ petrous part of the temporal,” is the bony capsule of the inner ear,
enclosing the labyrinth or essential organ of hearing, — in fact, it is
the skull of the ear, sometimes therefore called the otocrane — just as
ethmoidal parts form the “ skull of the nose,” and the sclerotal
bones represent a “skull of the eye.” The periotic consists of the
three bones already often mentioned, — the probtic, po, epiotic, ep,
and opisthotic, op, or anterior, superior, and posterior otocranial
bones, completely consolidated together, as well as with surround-
ing bones. The petrosal appears as an irregular^ protuberance in
the inner wall of the brain-cavity, at the lower back part. It seems
to be more extensive than it really is, because the great superior
semicircular canal, too large to be entirely accommodated in the
petrosal, has invaded the occipital bone, — the track of its bed in
that bone being sculptured in bas-relief (Fig. 70, asc). Behind this
semicircular trace, the deep groove of a venous sinus is engraved in
the bone, making the tract of the canal still more prominent (Fig.
70, sc). The top of the petrosal and contiguous occipital is the
door of a recess or fossa in which is lodged the great optic lobe
of the brain, partly divided from the general cavity for the cere-
bral hemisphere by a bony tentorium, like that which in mammals
separates the cerebellar from the cerebral fossse. On the vertical face
of the petrosal, or on the corresponding occipital surface, is a large
smooth-lipped orifice, at least of an inch in longest diameter ; it
leads to a tongue-like excavation of the bone, in which the, flocculus
of the cerebellum is lodged. In front, between the petrosal and
alisphenoid (or in the conjoined border of one or the other of these
bones) is a considerable foramen, conducting the second and third
divisions of cranial nerve 5 (see p. 261 ; Figs. 70, 71, ^) into the
o?^bit. Below the petrosal (in fact, between the opisthotic and the
exoccipital), near the border of the foramen magnum, is a foramen
(which may be subdivided into foramina), representing the foramen

278

GENERAL ORNITHOLOGY

PART II

lacerum posterius of mammals, transmitting cranial nerves 9, 10, 11
(see p. 262 ; Fig. 70, ®). The general space under description is
continued to the margin of the foramen magnum by the exoccipital
(Fig. 70, eo). Now on the vertical face of the petrosal itself — behind
foramen for 5, above that for 9, 10, 11, in front of the large floccular
orifice, will be seen a smooth-lipped depression, the meatus auditorius
internus (Fig. 7 0, ^), at the bottom of which are at least two separate
small foramina. A bristle passed in the upper (or anterior) one of
these two holes emerges outside the skull, in the tympanic cavity,
near the tympanic end of the Eustachian tube ; it has traversed the
interior of the petrosal, in a track known as the Fallopian nerviduct ;
it transmits cranial nerve 7 — the facial, or portio dura. A bristle
passed into the other of the two foramina may also be made to
come out in the tympanic cavity, but by a different track, for it
emerges through either the fenestra ovalis or the fenestra rotunda ;
it has traced the course of cranial nerve 8, — the auditory nerve, or
portio mollis. Both bristles have entered the common internal
auditory meatus, but the second one has traversed the ear-cavity
proper, through the lalyrinth of the ear, and come out at the tym-
panic vestibular orifice (fenestra ovalis), or at the tympanic cochlear
orifice (fenestra rotunda). Either passage is easily made, without
breaking down or indeed meeting with any bony obstacle, which
would not be the case with a mammal. Cranial nerves 7 and 8
were formerly counted as one (seventh) ; hence the name portio dura
“ hard portion ”) for the former, portio mollis (“ soft portion ”) for
the latter. The former, as said, traverses the petrosal bone and
escapes upon the face ; the latter, which is the true acoustic nerve,
or nerve of hearing, remains in the bone, being expended upon the
labyrinthine structures within — the vestibule, semicircular canals, and
cochlea, which constitute the walls of the cavities in which the essen-
tial organ of hearing is snugly encased.

If now, with a very fine saw — the saws now so much used for
fancy scroll-work will answer the purpose — the whole periotic mass
be cut away from the skull, and then divided in any direction, the
labyrinth can be studied. It is best to make the section in some
definite plane with reference to the axes of the whole skull, — the
vertical longitudinal, or vertical transverse, or horizontal, — as the
direction and relations of the contained structures are then more
easily made out. Four or five parallel cuts will make as many thin
flat slices of bone, affording eight or ten surfaces for examination ;
the whole course of the labyrinthine cavity can be seen in sections
which, when put together in the mind’s eye, or held a little apart
in their proper relations and visibly threaded with bristles, afford
the required picture very nicely. It is extremely difficult to chisel
out the affair from the bone in which it is embedded. At first

SEC. IV

ANATOMY OF BIRDS

279

glance the slices show a bewildering maze, — a continuous network
or lattice-work of bone, in which the unaccustomed eye will recog-
nise nothing but confusion. All this cancellated structure, however,
is pneumatic — the open-work tissue of the bone, containing air
derived from the tympanic or Eustachian cavities, and having nothing
to do with the ear-passages proper. Parts of the bony labyrinth
will soon be recognised by their firm smooth walls and definite
courses, as distinguished from the irregular interstices of the pneu-
matic bone -tissue. The bony labyrinth consists of an irregular
central cavity, the vestibule ; of a cavity, projecting like a beak
downward and backward from the vestibule, the cochlea ; and of three
horseshoe-shaped tubular cavities, above, behind, and below the
vestibule, the semicircular canals, the ends of whose hollows all open
into the vestibule. Imagine three hollow horse-shoes, with their ends
melted into a hollow inflation (vestibule), the opposite wall of which
is a hollow projection (cochlea) — or a hollow flat-iron (vestibule) with
a long nose (cochlea) and three hollow handles (the canals). Or,
see Figs. 84 to 87, representing the contained membranous labyrinth,
to which the containing bony labyrinth very closely conforms, as it
is simply the bony cavity whose walls encase the membranous and
other soft structures. According as the sections have been made,
numerous cross-cuts of the canals will be seen here and there as
circular orifices ; the canals themselves lying curled like worms in
the petrosal and occipital substance, their ends finally converging
to the vestibular cavity. As compared with those of man, the parts
are of great size ; in the eagle, the whole affair is as large as that
part of one’s thumb covered by the nail ; the whole length of the
superior semicircular canal is an inch or more j its calibre, I should
judge, being absolutely about as great as in man. The cochlea,
however, though not diminutive comparatively, is in a rudimentary
condition as far as complexity of structure is concerned, in all
Sauropsida, representing only the beginning of the cochlear struc-
ture of mammals. In the latter class, the cochlea is spirally coiled
or whorled on itself like a snail-shell (whence the name — cochlea, a
snail), making at least one turn and a half, sometimes five (two and
a half in man) ; with a centre-post or modiolus around which winds
a bony flange, the lamina spiralis, a membranous extension of which
to the cochlear out-wall divides the cavity into two compartments
or scalcB {scala, a flight of stairs) ; it is just like a spiral stairway,
only an inclined plane instead of a series of steps. The membran-
ous extension of the bony spiral lamina to the side-wall obviously
throws the cavity, as just said, into two spirals, which only inter-
communicate at the top, where the modiolus ends in a funnel-shaped
expansion, the infundibulum, beneath the apex of the snail-shell, the
cupola. A marble rolling down the upper stairway would fall into

28o

GENERAL ORNITHOLOGY

PART II

the vestibular cavity ; this division of the cochlea is therefore the
scala vestihuli. The marble starting from the other side of the in-

fundibulum would roll along the under stairway, and if nothing
stopped the way, would fall through the fenestra rotunda into the

SEC. IV

AJVA TOMY OF BIRDS

281

tympanic cavity ; this is therefore the scala tympani. The first
marble would also eventually reach the tympanum, through the
vestibule, and out of the fenestra ovalis, if the foot of the stapes were
unstepped (in life, of course, both these “ windows ” are closed by
membranous curtains). Now in birds the cochlear cavity and its
bony or cartilaginous contents are only the beginnings of such
structure — a strap-shaped or tongue-like protrusion from the vesti-
bule, as if a part of the first mammalian whorl, and very incom-
pletely divided into scala vestibuli and scala tympani by a gristly
structure (representing the modiolus and its lamina), which proceeds
from the bony bar or bridge between fenestra ovalis and fenestra
rotunda. (See Figs. 84, 85.) This structure is the most intimate
and essential part of the organ of hearing, for upon it spread the
terminal filaments of the auditory nerve. A human or any well-
developed mammalian cochlea is a thing of marvellous beauty, even
as to its bony shell — there is nothing to compare with its exquisite
symmetry ; while the spiral radiation of the nervous tissue intro-
duces yet other and more wondrous “ curves of beauty.”

The vestibule hardly requires special description ; it is simply
the central chamber common to the cochlear and canalicular cavities ;
receiving the mouth of the scala vestibuli of the cochlea ; the several
mouths of the separate or uniting semicircular canals ; opening into
tympanum by fenestra ovalis ; conducting to meatus auditorius
internus by the course of the auditory nerve. In the eagle, if its
irregularities of contour were smoothed out, it would about hold a
pea.

In the language of human anatomy, the three semicircular canals
are the {a) anterior or superior vertical, the (b) posterior or inferior
vertical, and the (c) external or horizontal ; and the planes of their
respective loops are approximately mutually perpendicular, in the
three planes of any cubical figure. In birds these terms do not apply
so well to the situation of the canals with reference to the axis of the
body, nor to the direction of the loops ; neither is mutual perpendicu-
larity so nearly exhibited. The whole set is tilted over backward to
some degree so that the {a) “ anterior ” (though still superior) loops
back beyond either of the others ; the (b) “ posterior ” loops behind
and below the (c) horizontal, which tilts down backward ; the verti-
cality of the planes of {a) and (5) is better kept. The canals may be
better known as the {a) superior (vertical), and (b) inferior (vertical),
and (c) internal (horizontal). Whatever its inclination backward,
there is no mistaking {a\ much the longest of the three, looping high
up over the rest, exceeding the petrosal and bedded in the occipital,
the upper limb and loop of the arch bas-relieved upon the inner
surface of the skull (Fig. 70, asc). It makes much more than a
semicircle — rather a horse-shoe. The inferior vertical (5) loops

282

GENERAL ORNITHOLOGY

PART II

lowest of all, though little if any of it reaches farther backward than
the great loop of {a) ; it is the second in size ; in shape it is quite
circular, — rather more than a half-circle. Its upper limb joins the
lower limb of (a), as in man, and the two open by one orifice in the
vestibule \ but it is not simj)le union, for the two limbs, before
forming a common tube, twine half-round each other (like two
fingers of one hand crossed). The loop of (^) reaches very near the
back of the skull (outside). The canal (c) is the smallest, and, as it
were, set within the loop of (5), though its plane is nearly the oppo-
site of the plane of (&) ; and the cavities of (5) and (c) intercom-
municate at or near the point of their greatest convexity, farthest
from the vestibule. This decussation of (6) and (c), like the twin-
ing inosculation of («) and (5), is well known. It may not be so
generally understood that there is (in the eagle if not in birds
generally) a third extra- vestibular communication of the canals. My
sections show this perfectly. The great loop of (a), sweeping past
the decussating-place of (&) and (c), is thrown into a cavity common
to all three. Bristles threaded either way through each of the three
canals can all three be seen in contact, crossing each other through
this curious extra-vestibular chamber, which may be named the
trivium, or “ three-way ” place. (The arrangement I make out does
not agree well with the figure of the owl’s labyrinth given by
Owen, Anat. Vert. ii. 134. The trivium is at the place where, in Fig.
84 or 85, the three membranous canals cross one another. It does
not follow, however, that these contained membranous canals inter-
communicate, and it appears from Ibsen’s figures that they do not.
Study of these admirable illustrations, with the explanations given
under them, should make the details perfectly clear to the reader.)

All that precedes relates to the bony labyrinth, — the scrolled
cavity of the periotic bone. The membranous labyrinth is a sac lying
loosely in the hollow of the bone, and shaped just like it, lining the
hollow of the vestibule and tubes of the semicircular canals. With-
drawn intact, it would be a perfect “ cast ” of the labyrinth. Ori-
ginally this sac is also continuous with one in the cavity of the
cochlea, called the membranous cochlea, which afterward becomes
shut off from the main sac. This shut-off cochlear part lies between
the scala tympani below and the scala vestibuli above ; its interior
is the scala media. If demonstrable in birds, it must be quite as
rudimentary as the other scalse. The membrane is not attached to
the bony walls of the labyrinth, but is separated by a space con-
taining fluid, the perilymph, which also occupies the scala vesti-
buli and scala tympani. A similar fluid, the endolymph, is contained
in the cavity of the membranous labyrinth, and scala media of the
cochlea ; in it are found concretions, or otoliths, of the same charac-
ter as the great “ ear-stones ” so conspicuous in many fishes. This

SEC. IV

ANATOMY OF BIRDS

283

lymph has a wonderful office — that of equilihmtion, enabling the
animal to preserve its equilibrium. The labyrinth and its contained
fluid may be likened to the glass tubes filled with water and a
bubble of air, by a combination of which a surveyor, for example,
is enabled to adjust his theodolite true to the horizontal. Some-
how a bird knows how the fluid stands in the self-registering level-
ling-tubes, and adjusts itself accordingly. Observations made on
pigeons show that “ when the membranous canals are divided, very
remarkable disturbances of equilibrium ensue, which very in char-
acter according to the seat of the lesion. When the horizontal
canals are divided rapid movements of the head from side to
side, in a horizontal plane, take place, along with oscillation of
the eyeballs, and the animal tends to spin round on a vertical
axis. When the posterior or inferior vertical canals are divided,
the head is moved rapidly backwards and forwards, and the
animal tends to execute a backward somersault, head over heels.
When the superior vertical canals are divided, the head is moved
rapidly forwards and backwards, and the animal tends to execute
a forward somersault, heels over head. Combined section of the
various canals causes the most bizarre contortions of the head and
body.” (Ferrier, Fund, of the Brain, 1876, p. 57.) Injury of the
canals does not cause loss of hearing, nor does loss of equilibrium
follow destruction of the cochlea. Two diverse though intimately
connected functions are thus presided over by the acoustic nerve,
— audition and equilibration.

Senses of Taste and Touch : Gustation and Taction. — The
hands of birds being hidden in the feathers which envelop the
whole body — their feet and lips, and usually much if not all of the
tongue, being sheathed in horn, these faculties would appear to be
enjoyed in but small degree. While it is difficult to judge how
much appreciation of the sapid qualities of substances birds may be
capable of, we must not be hasty in supposing their sense of taste
to be much abrogated. One who has had the toothache, or teeth
“ set on edge ” by acids, or painfully affected by hot or cold drinks,
may judge how sensitive to impressions an extremely dense tissue
can be. Persons of defective hearing may be assisted to a kind of
audition by an instrument applied to the teeth ; and it is not easy
to define the ways in which sensory functions may be vicariously
performed or replaced. Birds are circumspect and discriminative,
even dainty, in their choice of food, in which they are doubtless
guided to some extent by the gustatory sensations they experience.
As, however, only some human beings make these an end instead
of a natural and proper means to an end, the selection of food by
birds may be chiefly upon intuitions of what is wholesome. Such
purely gustatory sense as they possess is presided over by the

284

GENERAL ORNITHOLOGY

PART II

branches of the glossopharyngeal nerve which go to the back part of
the tongue and mouth. Though the chorda tympani nerve exists,
there is no lingual (gustatory) branch of the third division of the
fifth cranial nerve. Yet the latter, which goes in mammals to the
anterior part of the tongue, is less effectually gustatory than the
glossopharyngeal; as we know by the fact that the sensation of
taste is not completely experienced until the sapid substance passes
to the back of the mouth. Gustation is likewise connected with
olfaction ; the full effect of nauseous substances, for example, being
not realised if the nose is held. From these alternative considera-
tions, each one may estimate for himself how much birds know of
sapidity ; remembering also, how soft, thick, and fleshy are the
tongue and associate parts in some birds, as parrots and ducks, in
comparison with birds whose mouths are quite horny.

The beak is doubtless the principal tactile instrument ; nor does
its hardness in most birds preclude great sensitiveness ; as witness
the case of the teeth, above instanced. Sensation is here governed
by the branches of the fifth nerve. In some birds, in which also
the terminal filaments of this nerve are largest and most numerous,
the bill acquires exquisite sensibility. Such is its state in the snipe
family, in most members of which, as the woodcock, true snipe, and
sandpipers, the bill is a very delicate nervous probe. The Apteryx
also feels in the mud for its food, enjoying moreover the unusual
privilege of having its nose at the end of its long exploration.
Ducks dabble in the water to sift out proper food between the
“ strainers ” with which the sides of their beaks are provided ; and
the ends of the maxillary and mandibular bones themselves are full
of holes, indicating the abundance of the nervous supply (Fig. 63).

The senses of birds and other animals are commonly reckoned
as five — a number which may be defensively increased — as by a
sixth, the muscular sense, w'hich gives consciousness of strain or
resistance, apart from purely tactile impressions ; and perhaps a
seventh, the faculty of equilibration, which has a physical mechan-
ism of its own, at least as distinct and complete as that of hearing.
The ordinary “ five senses ” are curiously graded. Taction connotes
qualities of matter in bulk, as density, roughness, temperature, etc.
Gustation, matter dissolved in water — fluidic. Olfaction, matter
diffused in air — aeriformed. Audition, atmospheric air in undula-
tion. Vision, an ethereal substance in undulation. All animals
are probably also susceptible of hiogenation, which is the affection
resulting from the influence of biogen ; a substance consisting of
self-conscious force in combination with the minimum of matter
required for its manifestation.^

1 The reader who may be interested to inquire further in this direction is referred
to a publication entitled ; — Biogen : A Sjoeculation on the Origin and Nature of Life.

SEC. IV

ANATOMY OF BIRDS

285

c. Myology : the Muscular System

Muscular Tissue consists of more or fewer amoebiform animals j
separate colonies of which creatures, isolated in various parts of
the body, compose the individual different muscles. They are en-
veloped in fibrous tissue, the sheets of which are called fascice, and
the ends of which, usually attached to bones by direct continuity with
the periosteal covering of the latter, form tendons and ligaments.
The muscle - animals belong to a genus which may be termed
Myamceha, differing from other genera of the amoebiforms which
compose the body of a bird less in their physical character of being
elongated and spindle-shaped, or even filiform, than in their physio-
logical character of contractility. Under appropriate stimulus, as
the passage of a current of electricity, or the wave of biogen-
substance which constitutes a “nerve-impulse,” Myamocbce shorten
and thicken, tending towards a state of tonic contraction which, if
completed and long sustained, would cause them to become encysted
as spherical bodies ; but extreme contraction is never long con-
tinued. By alternate contraction and relaxation all the motions of
the body in bulk are effected. The capacity of, or tendency to, con-
traction is called the tonicity of muscular fibre. The simultaneous
contraction of any colony of Myamocbcc pulls upon the attachment of
the muscle at each of its ends ; in some cases approximating both
ends ; oftener moving the part to which one end is attached, the
other being fixed. The action of a muscle is upon the simplest
mechanical principles, — nothing more or less than pulling upon a
part, as by a rope, the line of traction being exactly in the line of
contraction of the muscle ; though it is often ingeniously changed
by the passage of tendons around a corner of bone, or through a
loop of fibrous tissue, as if through a pulley. Such movements as
those of a turtle protruding its head, or a bird thrusting its beak
forward, where muscle seems to 'jyus\ are fallacious ; when analysed,
the motion is invariably resolved into simple pulling. The swelling
up of a muscle in contracting must indeed impinge upon neighbour-
ing parts and shove them aside ; but that is an extrinsic result.
Muscles contract most powerfully under resistance to their tur-
gescence ; what is effected by the fasciae which bind them down ; —
what the athlete seeks to increase by bandaging his swelling biceps.
There are two species of Myamoeba. M. striata is the ordinary
striped fibre of voluntary motion, and also of some motion not

Abridged from a paper on the “ Possibilities of Protoplasm,” read before the Philo-
sophical Society of Washington, 6th May 1882. By Dr. Elliott Cones, etc. Washing-
ton, Judd and Detweiler. 8vo, 27 pp. Fifth edition. Boston, Estes and Lauriat,
1887.

286

GENERAL ORNITHOLOGY

PART II

under control of the will, as that of the heart. This species is
usually of a rich red colour (pale pink in many birds of the grouse
family), and is the ordinary “flesh” of the body. The other
species, ilf. Imis^ composes the pale or colourless smooth fibre of
the involuntary muscles, as those of the intestines, the gullet, etc.
A species of contractile tissue commonly referred to the genus
Desmamoebob (indifferent connective-tissue cells) is very near My-
amoeha Imvis ; example, mammalian dartos. The movements of
erectile organs, as the neat combs over the eyes of grouse, or the
turkey’s caruncles, are not in any sense myamoebic, but depend
mechanically upon influx of blood.

The Muscular System of Aves can only be touched upon ; it is
impossible in my limits to even name all the muscles, much less
describe them. I can only note the leading peculiarities, and pre-
sent a figure in which the principal muscles are named.

The subcutaneous sheet of muscle (of which the human “ muscles
of expression ” and platysma myoides are segregations) is broken up
in birds into a countless number of little slips which agitate the
feathers collectively, and especially the great quills of the wings
and tail. There are estimated to be 12,000 in a goose. The
prime peculiarity of birds’ musculation is the enormous develop-
ment of the pectorales, or breast muscles, which operate the wings.
The great pectoral, p. major or p. primus, arises from the sternal
keel, when that special bony septum between the fellow-pectorals
exists, and from more or less of the body of the sternum, passing
directly to the great pectoral or outer ridge of the humerus, near
the upper end of that bone. Its origin may even exceed the limits
of the sternum, invading the clavicle, etc. ; it may unite with its
fellow. It is the depressor of the humerus, giving the downward
stroke of the wing. The next pectoral, p. secundus or p. medius,
arises from much or most of the sternum not occupied by the first,
under cover of which it lies ; it passes also the humerus, but by an
interesting way it has of running through a pulley at the shoulder
it elevates that bone, giving the upward wing-stroke. A third
pectoral, p. tertius or p. minimus, arising from sternum, and often
contiguous parts of the coracoid bone, passes directly to the hume-
rus, supplementing the action of the first. A fourth muscle in
many birds acts upon the humerus from the sternum or coracoid,
particularly the latter. These four differ greatly in their relative
development. Such extent of the sternum and pectoral muscles
correspondingly reduces that of the belly-walls, and the abdominal
muscles are consequently scanty. Fixity of the spinal column in
the dorsal region diminishes the musculation of that part, the spinal
muscles being much better developed in the cervical region ; where,
in cases of some of the long-necked birds, there are curious con-

SEC. IV

ANATOMY OF BIRDS

287

trivances for the mechanical advantage of the muscle in flexing and
extending this mobile part of the body. Muscles of the hyoidean
apparatus acquire a singular development in woodpeckers. The
lower jaw is depressed particularly by muscle inserted into the end
of the mandible ; the upper is elevated by particular muscles operat-
ing the pterygoid and quadrate bones. Temporal, masseteric, and
ordinary pterygoid muscles close the jaws. They are unsymmetrical
in Loxia.

The diaphragm, the musculo-membranous partition which in
mammals divides the thoracic from the abdominal cavity, is only
represented in birds in a rudimentary condition. Macgillivray has
figured that of the rook as consisting of three fleshy slips, v, v, v,
passing from as many ribs, 4, 5, 6, to the pleural sac of the lungs,
t, t, in Fig. 101. It is best developed in the Apteryx.

The remarkable specialisation of both limbs, — the former for
flight, the latter for the perfectly bipedal locomotion which only
birds besides man enjoy, — results in corresponding peculiarities of
the muscular mechanism. Muscles beyond the shoulder are greatly
reduced in number and complexity from an ordinary quadrupedal
standard ; those of the legs are rather increased, and their configura-
tion, relative size, and to some extent their relations, are so much
changed, that great difficulty is experienced in identifying them
with the corresponding muscles of quadrupeds. The result is
great confusion in their nomenclature, which is still shifting, though
much has been done of late to give it precision. Attention has
recently been called by Garrod to the classificatory value of certain
muscles of the limbs. The tensor patagii, that muscle or those
muscles which may have elastic tendons, and by which the folds of
skin in the angles of the wing-bones are regulated, may have
different characters in different groups of birds. It has long been
known that particular muscles of the hind limb are in direct and
important relation to the prehensile power of the toes, and conse-
quently co-ordinated with the insessorial or the reverse character of
the foot. In the highest birds, Fasseres, the foot grasps with great
facility, owing to the distinctness or individuality of the flexor longus
kallucis, or bender of the hind toe. The ambiens (Lat. ambiens,
going around) is a muscle of which Garrod has even made so much
as to divide all birds into two primary groups according to whether
they possess it or not. The ambiens arises from the pelvis about
the acetabulum, and passes along the inner side of the thigh ; its
tendon runs over the convexity of the knee to the outer side, and
ends by connecting with the flexor digitorum perforatus — one of the
muscles which bend the toes collectively. When this arrangement
obtains, the result is that when a bird goes to roost, and squats on
its perch, the toes automatically clasp the perch by the strain upon

SEC. IV

ANATOMY OF BIRDS

289

the ambiens that ensues as soon as the leg is bent upon the thigh,
and the tarsus upon the leg, the weight of the bird thus holding it
fast upon its perch. The effect is as if an elastic cord were tied to
the hip joint, thence directed over the front of the knee and back
of the heel and so on to the ends of the toes. Obviously, such a
cord would be strained when the limb is bent, relaxed when the
limb is straightened out. The reader may observe a corresponding
effect of the muscular arrangement of his forearm by throwing the
hand as far back as possible ; the fingers tend to close by the strain
on the flexors in passing over what is a convexity of the wrist when
the hand is in that position. Passeres have no ambiens, the perfec-
tion of their feet in other respects answering all purposes. Birds
having it are termed homalogonatous or “normally-kneed” (Gr.
6/xaAos, hoTualoSj from 6/xos, homos, like, even, etc. ; yow, yovaros,
gonu, gonatos, knee) ; those wanting it are called anomalogonatous,
“abnormally-kneed.” The distinction prevails with much applica-
bility to various large groups of birds, and does good duty in
diagnosis when duly connected with other characters ; but surely
should not give name to primary groups founded upon it ! Other
muscles of the leg much used by the same sagacious and zealous
anatomist are the femorocaudal, accessory femorocaudal, semitendi-
nosus, and accessory semitendinosus. The whole five of these muscles
“ vary ; any one or more than one may be absent in different birds ;

. . . the constancy of the peculiarities in the different individuals
of each species, or the species of each genus, and very generally in
the genera of each family, makes it evident to any one working at
the subject that much respecting the affinities of the different
families of birds is to be learnt from the study of their myology, in
connection with the peculiarities of their other soft parts ; and that
these features will, in the long run, lead to a more correct classifica-
tion than one based on the skeleton alone, becomes almost equally
certain.” (Garrod, P. Z. S., 1873, p. 630.)

d. AnGIOLOGY : THE VASCULAR OR CIRCULATORY SYSTEMS

Blood and Lymph are the two media by the circulation of
which throughout the body the various amoeboid animals which
compose the tissues are fed, their waste repaired, and their dead
parts removed. Each species of Amoeba has the faculty of selecting
from the constituents of blood and lymph its appropriate food ; and
of converting such nourishment into its own proper substance.
Eefuse matters are either drained off by the kidneys and voided as
excrement, or swept by the current of blood into the lungs and
there cremated. The stream of lymph is a feeder to the blood, and
when the mingled currents are no longer distinguishable has become
u

290

GENERAL ORNITHOLOGY

PART II

blood. The machinery of circulation is two sets of vessels — the
hcematic, or vascular system proper, consisting of the heart, arteries,
veins, and capillaries, for the blood-circulation ; and the lymphatic^
consisting of lymph-hearts and vessels, for the flow of lymph. The
lymphatics, converging from all parts of the body, and especially
from the intestines, end in vessels which pour the lymph into the
veins of the neck. The heart is the central organ of the blood-
circulation, by which that fluid is pumped into all parts of the body
through the arteries or efferent vessels ; straining through the net-
work of capillaries, it returns to the heart through the veins, or
afferent vessels. The set of efferent vessels is the arterial system ;
that of afferent vessels is the venous system. The blood in arteries
excepting the pulmonary is bright red ; that in veins excepting the
pulmonary is dark red. The change from bright to dark occurs in
the capillaries of the system at large ; the change from dark to
bright only in the capillaries of the lungs and air-sacs. The systemic
blood circulation is completely separated from the pulmonic in all
animals in which, as in birds, the right and left sides of the heart
are separated from each other ; such circulation is said to be double ;
that is, arterial and venous blood only mingles in the capillaries,
whether of the lungs or others, and therefore at the periphery of the
vascular system : the heart being the centre of that system. Blood,
in all or some of its constituents, permeates absolutely every tissue
of the body. Those tissues whose capillaries are large enough for
the passage of all the constituents of blood are said to be vascular ;
those which only feed by sucking up certain constituents of the
blood, and have no demonstrable capillaries, are called non-vascular.
But nutrient fluid penetrates the densest tissue, as the dentine of
teeth ; no permanent tissues are really non-vascular, or they would
soon die, as do feathers, which require to be renewed once a year
or oftener.

Lymph and the lymphatics are noticed further on. Blood con-
sists of water in which several ingredients are dissolved, and cer-
tain solid bodies are suspended. Its water is salted, albuminated,
fibrinated, and corpusculated. The proportions, which vary in
different birds and at different times in the same bird, are in round
numbers: water 80, fibrine and corpuscles 15, albumen and salts
5 = 100 parts. Withdrawn from the body and allowed to settle,
blood separates into two parts, serum and coagulum. The serum is
the clear yellowish salty albuminous water ; the clot is the fibrine,
in the meshes of which are mired the corpuscles, reddening the
whole mass. The plasma, plasm or plastic material of the blood, is
its substance dissolved in water ; that is to say, minus the solid cor-
puscles. These latter interesting little bodies are a myriad of
minute animals, which swim in the life-current, and are named

SEC. IV

ANATOMY OF BIRDS

291

HcBmatammla cruentata. They have been supposed to be of two
species ; but the so-called white blood corpuscles, or leucocytes, in-
distinguishable from lymph corpuscles, are simply the formative
stages of the red blood-discs. In its early colourless stage, the
Hcematamoeba is a nucleated mass of protoplasm {protoplasm is the
indifferent substance out of which all animal tissue is derived), of no
determinate size or shape, exhibiting active amoeboid movements.
Later in the life of the minute creature, it passes into a sort of
encysted state, in which it reddens and acquires definite dimen-
sions and configuration. In birds these “blood-discs” are flat,
elliptical, and nucleated, that is, containing a kernel ; they average
in the long diameter in the short, g-gVoj ^n inch. Thus

they differ decidedly from the flat, circular, non - nucleated, red
blood-discs of Mammalia, which latter are supposed to be rather free
nuclei than perfected Hoematamoehce. The red colour of blood is en-
tirely due to the presence of these unicellular animals. The energy
of respiration, and corresponding activity of circulation in birds,
make them hcematothermal, or hot-blooded ; the pulse is quickest,
the blood hottest, and richest in organic matter, in these of all
animals.

The Heart is a hollow muscular organ, at the physiological
centre of the haematic vascular system. Its muscle presents the
principal exception to the rule, that the contractility of Myamceha
striata (see p. 285) is subject to voluntary control. It is the most
industrious organ of the body, never ceasing its rhythmic systole and
diastole, or contraction and dilatation, from the moment of the first
pulsation in the contractile vesicle which begins it, to that when
the “ muffled drum ” gives the last beat of the “ funeral march to
the grave.” The arteries are the elastic, thick-walled, branching
tubes which leave the heart on their way to the body at large ;
their pulsations, over which the vasomotor nervous system presides,
are isochronous with the heart-beats, and arterial blood thus flows
in jets. The veins are the vessels converging from all parts ; thin-
walled, less elastic, with more equable current. The capillaries are
the communicating vessels, of such size as just to permit the Haema-
tamoebas to pass through ; their network represents the termina-
tions of arteries and the commencements of veins. The heart in
adult birds is completely double ; i.e. the right and left sides are
perfectly separated. It is also completely four -chambered ; i.e.
there is an auricle and a ventricle on each side, which communicate ;
in embryonic life the two auricles communicate by the foramen ovale,
which afterward closes. Arteries proceed from the strong muscular
ventricles ; veins are received by the weaker auricles. The course
of the blood is : From the body, excepting the lungs, it comes dark
and heavy with products of decomposition, through the caval veins

2Q2

GENERAL ORNITHOLOGY

PART II

into the right auricle ; from right auricle through the auriculo-ven-
tricular opening into right ventricle ; from right ventricle through
the 'pulmonary arteries to the lungs ; in the capillaries of which it is
relieved of its burden. There decarbonised and oxygenised, the
bright red aerated blood returns through the pulmonary veins to the
left auricle ; through the corresponding auriculoTventricular opening
to the left ventricle, which pumps it out through the aorta and other
arteries to the capillaries, and so to the veins and heart again.
Thus the pulmonary arteries convey black blood, the pulmonary
veins red blood ; the reverse of the usual course. Before lungs come
into play, in the egg, the blood is purified in the allantois, an em-
bryonic organ which then sustains a respiratory function. Besides
the pulmonary there is another special circulatory arrangement, the
hepatic portal system of veins, by which blood coming from the
chylopoetic viscera (stomach, intestines, etc., which make chyle in
the process of digestion) strains through the liver before reaching
the heart. There is no renal portal system in birds.

The heart of birds is not peculiar in its conical shape, but is
more median in position than in mammals. There being no com-
pleted diaphragm, the pericardial sac which holds it is received in
a recess between lobes of the liver. The right ventricle is much
thinner- walled than the left ; the auricles have less of the elonga-
tion which has caused their name (“little ears” of the heart) in
mammals. The right auriculo- ventricular valve, which prevents
regurgitation of blood, instead of being thin and membranous, is a
thick fleshy flap which during the ventricular systole applies itself
closely to the walls of the cavity. The pulmonary artery and the
aorta are each provided at their origination with the ordinary three
crescentic or semilunar valves, as in mammals. The pulmonary
artery arises single, forking for each lung. The pulmonary veins
are two. The systemic veins, or vence cavce, bringing blood from the
body at large, are three — two precaval, from head and upper ex-
tremities, one postcaval, from trunk and lower extremities. The
aorta, almost immediately at the root of that great trunk, Figs. 90-
95, h, divides into three primary branches ; right, ri, and left, li,
innominate arteries, conveying blood to the neck, head, and upper
extremities j and main aortic, a, which curves over to the right (left
in mammals) and supplies the rest of the body. More precise state-
ment is, perhaps, that the aortic root, h, first gives off the left in-
nominate, li, then at once divides into right innominate, ri, and
main aortic trunk, a, (right). It represents the fourth primitive
aortic arch of the embryo. On the whole, the avian heart is a great
improvement on that of most reptiles, though nearly resembling
that of Crocodilia ; it is substantially as in any mammal, though
differing in its fleshy right auriculo-ventricular valve, two instead

SEC. IV

ANATOMY OF BIRDS

293

of one precaval vein, right instead of left aortic arch, and mode of
origin of the primary aortic branches.

The zoological interest of the avian blood-vessels centres in the
carotid arteries, which, with the vertebral arteries, supply the neck
and head. The carotids may be single or double ; and other details
of their disposition correspond well with certain families and orders
of birds. They are the first branches of the innominates. In most

Figs. 90-95. — Diagrams of carotid arteries of birds : h, root of aorta ; a, arch of aorta, to
the right side ; li, left innominate ; ri, right innominate ; Is, left subclavian ; rs, right sub-
clavian ; Ic, left carotid ; rc, right carotid. (1) Fig. 90. Aves Mcarotidince normales, with two
carotids, both alike. (2) Fig. 91. Aves Icevo-carotidinoe, with left carotid only. (3) Fig. 92.
Aves Mcarotidince cibnormales, certain parrots, with two carotids, not alike. (4, 5, 6) Aves
conjuncto-carotidince, with two carotids, which speedily unite in one. (4) Fig. 93, bittern,
both alike. (5) Fig. 94, flamingo, left very small. (6) Fig. 95, cockatoo, right very small.
(Copied by Shufeldt from Garrod.)

birds, there is but one carotid, the left ; in a few, one, formed by
early union of two ; in many, two, long distinct. The arrangement
will be perceived by the diagrams taken from Garrod’s admirable
paper (P. Z. S., 1873, p. 457). In nearly the words of this author :
1. In what may be termed the typical arrangement (though it is not
the usual one), two carotids, of equal size or nearly so, run up the
front of the neck, converging till they meet in the middle line,
and so continue up to the head, on the front of the bodies of the

294

GENERAL ORNITHOLOGY

PART II

cervical vertebrse, in the hypapophysial canal. Birds with this
arrangement Garrod calls aves Ucarotidinm normales (Fig. 90). 2. In

most birds the carotid branch of the right innominate being not
developed, only the left, of larger size, traverses the hypapophysial
canal ; but it bifurcates before reaching the head, thus producing
two carotids, distributed as if there had been two all the way up.
Such birds are said to have a left carotid, and are termed aves
Icevo-carotidinoe (Fig. 91). 3. In certain parrots only, with two

carotids, the right is as in (1), but the left runs superficially along
the neck with the jugular vein and pneumogastric nerve j such birds
are aves Ucarotidinoe abnormales (Fig. 92). 4. Two carotids, arising

normally, unite almost immediately, and the single trunk runs to
near the head, just as if there were two as in (1) ; then it bifurcates,
as in birds with left carotid only (2). Such birds are termed aves
conjuncto-carotidince. Special cases of (4) are : in the bittern, the
two roots are of nearly equal size (Fig. 93) j in the flamingo, the
left is very small (Fig. 94) ; in a cockatoo, the right is very small
(Fig. 95). Parrots display all four of the arrangements; the cases
of the bittern and flamingo are unique. The question is thus for
nearly all birds narrowed to whether there be two normal carotids
(1), or the left only (2). Observations upon three hundred genera
show two in one hundred and ninety-three, in one hundred and
seven the left only ; but the numerical proportion of Passerine genera
makes (2) the most frequent arrangement. There is but one carotid
in all Passeres as far as known ; in most Cyjpselidoe ; in Trogonidm,
Meropidce, Upupidce, Rliamphastidce, some Psittaci, the Turnicidce,
Megapodidce, Podicijpedidce, Alcidce, Piheidoe, Artery gidce. Thus in

Passeres, Columbm, Accipitres, Grallce, and Anseres the carotid arrange-
ment is an ordinal character, all but the first named of these great
groups having two. The character separates most of the families of
“Picarian ” birds, and also distinguishes the families Phoenicopteridoe,
Megapodidce, Cracidoe, Turnicidce, Podicipedidce, and family groups of
the Ratitce, from among one another. It is apparently only a
generic character in Psittaci, and in Cypselidce, Ardeidce, and Alcidce.

Beaching the skull, the carotids burrow in the bone, between
the basitemporal plate and the true floor of the skull, and enter the
cranial cavity by the “ sella turcica ” (the original pituitary space) ;
their anastomosis furnishes a sort of “circle of Willis.” (Figs. 66,
69, 70, ic.)

Both limbs of birds have a prime peculiarity of their arteries as
compared with mammals. In the fore limb, the blood supply being
chiefly absorbed by the immense pectoral muscles, vessels which in
mammals are small axillary branches appear like the main continua-
tion of the subclavian trunk, and the brachial arteries are correspond-
ingly reduced. In the leg, the main source of supply is the great

SEC. IV

ANATOMY OF BIRDS

295

ischiac artery, the femoral being small. This ischiac artery corre-
sponds to the twig which in man accompanies the great sciatic nerve
{comes nervi iscTiiatici) ; and the rare human anomaly of a posterior
main vessel of the thigh is therefore a reversion (atavism) to the
avian rule. There is no single proper renal artery to the kidney.

The Lymphatics of birds consist chiefly of a deep set accompany-
ing the main blood vessels, forming various plexus, — nodes, “ glands,”
or “ lymph hearts,” in their course. Superficial lymphatics, so promi-
nent in mammals, are little developed, though lymphatic glands are
found in the armpit and groin of some birds. These are the
systemic lymphatics ; a special set, the lacteals, arise by numberless
twigs in the course of the small intestine, uniting and reuniting to
form at length two (not one as in mammals) main tubes, which lie
along either side of the spinal column. These are the thoracic ducts ;
which terminal trunks of the whole lymphatic system empty into
the right and left jugular veins at the root of the neck. The contents
of the vessels differ correspondingly. Pure lymph is a pale, limpid,
albuminous fluid, containing when maturely elaborated a number of
irregular amoeboid bodies indistinguishable from the white formative
corpuscles of the blood (p. 291). It is strained out of the tissues at
large, being that material, not yet effete, which is still fit for feed-
ing the blood. The lacteals contain chyle, — the other kind of lymph,
drained off by the mucous membrane of the intestine from the j^re-
pared food in that tube ; an albuminous fluid, milky or cloudy from
the abundance of oil-globules, which, after mingling with the systemic
lymph, is poured directly into the current of the blood, in the
manner above said. Since the lacteals do not appear to begin with
open mouths, the chyle must soak into them through the lining
membrane of the intestines j and as this consists of a layer of amoeba-
like animals, through whose bodies the chyle passes, it is quite true
to say that the whole organism is nourished upon the excrement of
amoebas.

e. Pneumatology : the Eespiratory System

The Organs of Respiration provide for the ventilation of the
body. Since the respiratory process is also calorific, they likewise
furnish a heating apparatus. They consist essentially of air-passages
and air-spaces connected with lung-tissue, being therefore pulmonary
organs. No other animals are so thoroughly permeated as birds
with the atmospheric medium in which they live ; in no others are
the respiratory functions so energetic and effectual. The lung may
be likened to a blast-furnace for the combustion of decayed animal
matter ; purification of the blood and warming of the body being
two inseparable results obtained. Dark blood flowing to the lungs.

296

GENERAL ORNITHOLOGY

PART II

heavy with effete carbonaceous matters, is there relieved of its
burden and aerated by the action of oxygen ; the products of com-
bustion being exhaled in the form of carbonic dioxide and water.
Aside from the proper lung-tissue, the capillary substance of the
immense air-sacs tends to the same result. There is likewise, in
birds, a lesser system of ventilation, by which air is admitted to
cranial bones through the Eustachian tubes \ but this is unconnected
with the proper respiratory office. Pulmonary tissue consists chiefly
of a wonderful net (a rete mimUle) of capillaries, interlacing in every
direction, bound together and supported by fine connective tissue,
and invested with membrane so delicate that their walls seem naked,
their exposure to the air being thus very thorough. Air gains such
intimacy with the capillaries through the larynx, trachea (Fig. 101, (?),
and Ironchial tubes (r, r), these being the primary air-passages. But
all the bronchial tubes do not subdivide into the ultimate air-cells ;
some large ones run through the lung, pierce its surface (as at u, u,
Fig. 101), and end in that system of enormous air-spaces for which
the respiratory system of birds is so remarkably distinguished, —
like a heap of soap-bubbles, blown up en masse from a bowl of fluid ;
the extra-pulmonary air-spaces being the larger superficial bubbles, the
minute vesicles of lung- tissue proper being little bubbles just formed.
In this way air penetrates even the hollow skeleton of most birds.

The Lungs of Birds (Fig. 101, ^, ^), notwithstanding their heated
energy of respiration, are anatomically more like those of reptiles
than of mammals. They are not shut by a diaphragm in a special
division of the great thoracic-abdominal cavity of the body, but
extend from the apex of the chest as far as the kidneys, in the
pelvic region. They are not divided into lobes, as in mammals, nor
do they as in that class float freely in the chest by their mooring at
their roots ; nor, again, are they completely invested by a serous
membrane forming a closed pleural cavity. They are fixed in the
dorsal region of the general cavity, covered in front with pleura, with
which slips of the rudimentary diaphragm {v, v, v) are connected ;
but on the dorsal surface are accurately moulded to the intercostal
spaces, showing the impressions of the ribs and vertebrae, — just as
the lobulated kidneys are stamped with the sacral inequalities of
surface. They are, as usual, two, right and left ; their “ roots ” are
the bronchi {r, r), the pulmonary arteries and veins, nerves, and
connective tissue.

The Pneumatoeysts. — A bird is literally inflated with these
great membranous receptacles of air, and draws a remarkably “ long
breath,” — all through the trunk of the body, in several pretty
definite compartments ; in many, or most, or all, of the bones ; in
many intermuscular spaces ; in some birds also throughout the
cellular tissue immediately beneath the skin. These cysts vary so

SEC. IV

ANATOMY OF BIRDS

297

much in extent and disposition as to be not easily described
except either in the most general terms already used, or with
particularity of detail for different species. According to Owen,
however, the usual disposition is : An interclavicular air-space,
quite constant : this, with its cervical prolongations, furnishes the
great “ air-drums ” of the pinnated grouse and cock-of-the-plains.
Anterior thoracic, about the roots of the lungs. Lateral thoracic, pro-
longed to axillary, and to spaces and passages in the wings, includ-
ing the hollow humerus. Large hepatic or posterior thoracic, about
the lower part of the lung and the liver. Abdominal, right and left,
of great size, from the lower part of the lung where the longest
bronchial tubes open very freely j extending to pelvic and inguinal
compartments, whence femoral sacs, the hollow of the femur, etc.
The subcutaneous cells are enormously developed in the pelican and
gannet ; the extensive areolar tissue being thoroughly pneumatic,
and furnished with an arrangement of the cutaneous muscle (pan-
niculus carnosus) whereby, apparently, the air may be rapidly and
forcibly expelled by compression. A similar muscle develops in
some birds in connection with the interclavicular air-space. (The
pneumaticity of the skeleton has been already treated.)

The purpose of this extensive respiratory apparatus is thus
dwelt upon by the great English anatomist just cited : “ The
extension from the lungs of continuous air-receptacles throughout
the body is subservient to the function of respiration, not only by a
change in the blood of the pulmonary circulation effected by the air
of the receptacles on its repassage through the bronchial tubes ; but
also, and more especially, by the change which the blood undergoes
in the capillaries of the systemic circulation which are in contact
with the air-receptacles. The free outlet to the air by the bronchial
tubes does not, therefore, afford an argument against the use of the
air-cells as subsidiary respiratory organs, but rather supports that
opinion, since the inlet of atmospheric oxygenated air to be diffused
over the body must be equally free. A second use may be ascribed
to the air-cells as aiding mechanically the action of respiration in
birds. During the act of inspiration the sternum is depressed
[lowered from the back-bone in horizontal position of a bird], the
angle between the vertebral and sternal ribs made less acute, and
the thoracic cavity proportionally enlarged ; the air then rushes into
the lungs and thoracic receptacles, while those of the abdomen
become flaccid ; when the sternum is raised or approximated towards
the spine, part of the air is expelled from the lungs and thoracic
cells through the trachea, and part driven into the abdominal
receptacles, which are thus alternately enlarged and diminished with
those of the thorax. Hence the lungs, notwithstanding their fixed
condition, are subject to due compression through the medium of

298

GENERAL ORNITHOLOGY

PART II

the contiguous air-receptacles, and are affected equally and regularly
by every motion of the sternum and ribs. A third use, and per-
haps the one which is most closely related to the peculiar exigencies
of the bird, is that of rendering the whole body specifically lighter ;
this must necessarily follow from the desiccation of the marrow and
other fluids in those spaces which are occupied by the air-cells, and
by the rarefaction of the contained air from the heat of the body. . . .
A fourth use of the air-receptacles relates to the mechanical assistance
which they afford to the muscles of the wings. This was suggested
by observing that an inflation of the air-cells in the gigantic crane
{Ciconia argala) was followed by an extension of the wings, as the
air found its way along the brachial and antibrachial cells. In large
birds, therefore, which, like the argala [or like the wood-ibis, Tantalus
loculator], hover with a sailing motion for a long -continued period
in the upper regions of the air, the muscular exertion of keeping
the wings outstretched will be lessened by the tendency of the
distended air-cells to maintain that condition. It is not meant to
advance this as other than a secondary and probably partial service
of the air-cells. In the same light may be regarded the use as-
signed to them by Hunter, of contributing to sustain the song of
birds and to impart to it tone and strength. It is no argument
against this function that the air-cells exist in birds which are not
provided with the mechanism necessary to produce tuneful notes ;
since it was not pretended that this was the exclusive and only
office of the air-cells.” (Owen, Anat. Vert., ii. 1866, p. 216.)

Though nothing like them exists in mammals, it must not be
inferred that these air-pouches are unique in birds. The general
pulmonary mechanism is reptile-like, and the ornithic development is
simply a logical extreme of arrangements found in reptiles and lower
vertebrates, — even to the swim-bladder of a fish, which is morpho-
logically and homologically pulmonary, though fishes’ gills are
functionally, and therefore analogically, their lungs, i.e. their
respiratory apparatus.

The Trachea (Or. rpayeta, tracheia, rough) or “ asper-artery ”
answers perfectly to its English name, windpipe. It is the tube
which conveys air to and from the lungs (Fig. 101, \ 0 to ^). It
commences at the root of the tongue by a chink in the floor of the
mouth (Fig. 101, c), runs down the neck in front between the

gullet and the skin, and ends below by forking into right and left
bronchus (Fig. 101, \ r, r). It is composed of a series of very
numerous gristly or bony rings connected together by elastic
membrane. Lengthening and shortening, effected by muscles to be
presently noted, is permitted by a very ingenious and interesting
construction of these rings, which will be clearly understood with
the help of the figures (96, a, h, 97, \ \) borrowed from Macgillivray’s

SEC. IV

ANATOMY OF BIRDS

299

admirable account. When contracted, the rings look like an
alternating series of lateral half-hoops, as in Fig. 96, a; when
stretched to the utmost, as in Fig. 96, b, they are clearly seen to be
annular, or completely circular. The curious bevelling of the right
and left sides of each ring alternately is ^ ^

shown in Fig. 97, \ and Fig. 97,
represents the same two rings put to-
gether. The principle by which any
two rings slip partly over each other on
alternate sides is something like that
upon which a cooper fastens the ends of
any one barrel-hoop without any nailing
or tying. The rings are in some birds
perfectly cartilaginous : in most they
become osseous. The trachea is moved
by lateral muscles, which not only
shorten the tube by approximating the
rings, but also drag the whole structure
backward, by their attachment to the
clavicle and sternum. The strip, or two
strips, of muscle lying upon each side
of the trachea, is the contractor trachece
(Fig. 101, ss, ss) ; the most anterior,
when there are two, as soon as it
leaves the tube to go to the clavicle,
becomes the clidotrachealis, or clidohyoid, Fig. 101,
other is similarly the sternotrachealis.

Fig.

i. — a, an inch of trachea,
contracted to the utmost, the rings
looking like alternating half-rings ; &,
the same, stretched to two inches,
the rings evidently complete, with
intervening membrane. (After Mac-
gillivray.)

^ /, // the
The latter may be a
direct continuation of the contractor, as in Fig. 101, \ the loose
strips under q, or apparently arise separately from the side of the
lower end of the tube, as in Fig. 101, e. (Other muscles are to

be described with the
larynx superior and
inferior.) The trachea
is long in birds, pro-
portionate to the exten-
sion of the neck; it is

Fig. 97. — 1, 2, left-hand, two tracheal rings, separate, as Very fleXUOUS, follow-
in Fig. 96, 6; 1, 2, right hand, the same put together, as in witln pgqp fVip

Fig. 96, a. (After MLgiilivray.) ° ’ mg witii ease tue oenus

of the neck in which
it lies so loosely. Its cross section is oval or circular ; but all that
relates to the configuration and course of the pipe requires special
description, — so variable is the organ in different birds. It is

subject to dilatations and contractions in any part of its extent, and
to deviations from its usual direct course to the lungs. Minor
modifications must be passed over. The most remarkable expan-

300

GENERAL ORNITHOLOGY

PART II

sions of the lower part of the tube occur in many sea-ducks and
mergansers {Fuligulince, Merginm), and some other birds ; several
lower rings of the trachea being enormously enlarged and welded
together into a great bony and membranous box, of wholly irregular,
unsymmetrical contour. Such a structure, represented in Figs. 3
and 98, is termed a tracheal tymjpanum, or labyrinth. It is not a
part of the voice -organ proper, but may act as a reverberatory
chamber to increase the volume of the sound, without however
modulating it. Being chiefly developed in the male, it is a kind of
secondary sexual organ. The vagaries of the windpipe are still
more remarkable. Very generally, in cranes and swans, the trachea
enters the keel of the sternum, which is excavated to receive it,

and where it forms one or more
coils before emerging to pass to
the lungs. This curious winding
is carried to an extreme in Grus
americana, the whooping crane, in
which the windpipe is about as
long as the whole bird, and about
half of it — over two feet of it ! —
is coiled away in the breast-bone
(Fig. 99). The same thing occurs
in G. canadensis to a less extent
(Fig. 100). In a guinea-fowl.
Gutter acristata, a loop of the
trachea is received in a cup formed
by the apex of the clavicles. In
various birds, as some of the
.xia.c V.. curRssows (Cracidm), the caper-

seen from behind, nat. size. Dr. R. W. Shu- cailHe (TetraO uroqallus), a gOOSe,

Anseranas sermpatmata, and the
female of the curious snipe, Rhynchcea australis, the trachea folds
between the pectoral muscles and the skin.

The Larynx (the Gr. name, Xdpvy^, larugx) is the peculiarly
modified upper end of the trachea (Fig. 101, \ and ^ to ^^). In
mammals it is a complicated voice-organ, containing the vocal chords
and other consonantal apparatus ; in birds the construction is
simpler, as the larynx merely modulates the sound already produced
in the lower end of the tube. It lies in the floor of the mouth, at
the root of the tongue, between the forks of the hyoid bone, resting
upon the urohyal. Besides its attachments of mucous and other
membrane, it is connected with the hyoid bone by a pair of thyro-
hyoid muscles (V/)? usually with the rest of the trachea by
prolongations of the sterno- and clido-tracheales. It is usually a
small, simple, conical “ mouthpiece ” of the pipe (^, a), without the

SEC. IV

ANATOMY OF BIRDS

301

dilatation which renders the corresponding structure — the “ Adam’s
apple,” — so conspicuous in the human throat. Below, it communi-
cates directly with the pipe : above, it opens into the mouth by the
glottidean fissure^ or rima gloUidis (^, c), a median lengthwise chink,
which opens and shuts as its sides diverge or close together, and
which is further defended in front by a folding of the mucous
membrane of the mouth, constituting a rudiment of that curious
trap-door arrangement which, when fully developed, is called the
epiglottis (^, d, e). Exclusive of two Irohen upper rings of the
trachea (®, g), the cartilages (or oftener bones, — for they generally
ossify) of the larynx are five. One is a large single median and
inferior piece, the thyroid, or shield-piece (\ a), forming the

most substantial part of the structure. It is somewhat triangular
or oblong, running to an obtuse end in front ; and with sides and
posterior angles which curl upward behind. To its lateral posterior
corner is attached on each side the small “horn” or corniculum
laryngis (^, V). There is a small median upper posterior piece,

supposed to represent all there is of the cricoid (^, c), which in

man makes a ring around the larynx below the thyroid. To the
cricoid, as to a base, are attached a pair of straight slender arytenoids
(®, d), projecting forward along the upper surface of the larynx :

these form the rima glottidis, — the fissure of the glottis being
between them. The arytenoids are attached in front by slender
ligaments to the end of the thyroid (^, the little slips between d and
e), and they are supplemented by cartilaginous edges (®, /, /) ; but
there are no true vocal chords. Besides the extrinsic thyrohyoid
muscles, which pass from the larynx to the tongue -bone, the
laryngeal parts are operated by intrinsic muscles, the sum of the
motion given by which is the opening and shutting of the glottis
by drawing apart or pulling together the arytenoids. Four pairs
of such muscles are described for some birds. As named and
figured by Macgillivray for the rook, there are : the thyroarytenoids,
which are the openers of the glottis (^, ; the oblique arytenoids

(10, 3,3^ . thyrocricoids (^^, ; and the posterior thyrocricoids,

(^^ and

The Syrinx (Gr. o-vptyf, surigx, a pipe) or Lower Larynx is the
voice- organ of birds ; in most respects a more complicated structure
than the larynx proper, and one so differently constructed in
different birds that it affords characters of great significance in
classification. The highest group of Passeres, for example, is
signalised by the elaboration of this musical organ, the marvellously
adroit fingering of the keys of which by the little muscular
performers sends through the tracheal sounding-pipe the tuneful
messages of bird’s highest estate. A few degraded or disgraced
birds, as the ostrich and the American vultures, have no bucolic

302

GENERAL ORNITHOLOGY

PART II

organ at all, the trachea forking as simply as possible. Others, as
the common fowl, have a fair syrinx, but no muscles whatever to
modulate their pastoral lays. Others have one, two, or three pairs
of intrinsic muscles ; to which may or may not be added a sterno-
tracheal with syringeal attachment. It is not so much the bulk or
mere fleshiness of the syrinx that indicates musical ability ; but the
distinctness of the several muscles, and the mode of their insertion.

Fig. 99. — Coiling of the windpipe in the sternum of Grns americana ; reduced.

which result in endless combinations of rotating and rocking move-
ments of the parts, whereby an infinite modulation of the musical
tones becomes possible. In Oscines there are normally five or six
pairs of muscles, without counting the extrinsic sternotracheales ;
and the gist of the arrangement, in these melodious Passeres, is the
attachment of the muscles to the ends of the upper bronchial half-
rings, as far as the third one. As Professor Owen remarks with
appreciative feeling, “ the manifold ways in which the several parts
of the complex vocal organ in Cantores may be affected, each of the

SEC. IV

ANATOMY OF BIRDS

303

principal bony half-rings, as one or the other end may he pulled,
being made to perform a slight rotatory motion, are incalculable ;
but their effects are delightfully appreciable by the rapt listener to
the singularly varied kind and quality of notes trilled forth in the
stillness of gloom by the nightingale.”

I should be able to make the plan of the syrinx clear to the
student with the assistance of Macgillivray’s beautiful figures.
These are drawn from the rook, — a corvine croaker, indeed, but
one whose syrinx is in good order, though he has never learned to
play. As the modifications affect principally the soft parts covering

and moving the music-box, one description of the latter is applicable
to most birds. The last lower ring, or piece composed of several
fused rings, of the trachea, at its bifurcation into bronchi, is enlarged
or otherwise modified (Fig. 101, ahd)^ and crossed below from
front to back by a bony bar, the pessulus at b ; a)^ or bolt-bar,

which, dividing it into lateral halves (as at forms thus two
lateral openings instead of one median tube, — the beginnings of
each bronchial tube. A membranous plate, strengthened by
cartilage, rises vertically into the tracheal tube, forming a septum, or
median partition, between the orifices of each bronchus. The free
curved upper margin of this septum, extending from front to back

SEC. IV

ANATOMY OF BIRDS

305

bird ; nat. size, after Macgillivray. 1. a, tongue ; 6, basibranchial, commonly called urobyal ;

c, c, horns of hyoid bone ; d, d, geniohyoid muscles ; e, e, stylohyoid muscles ; f, f, clidohyoid
muscles ; g, h, i, oesophagus ; j, proventriculus, or secretory stomach ; 7c, gizzard, or gigerium,
the muscular stomach ; I, m, n, n, intestine, duodenum to rectum ; 0, p, trachea, or windpipe ;
q, inferior larynx, or syrinx ; r, r, right and left bronchus ; ss, ss, contractor muscles of trachea ;
t, t, lungs, with u, u, apertures communicating with thoracic air-cells ; v, v, v, three pairs of
muscular slips answering to a rudimentary diaphragm ; 1,2, 3,4,5, 6,7, as many ribs.— 2. Hyoid
bone ; a, glossohyal, tipped with cartilage, its posterior horns being ceratohyals proper ; b,
basihyal ; c, basibranchial proper, commonly called urohyal ; d, d, ceratobranchials proper,
commonly called apohyals ; e, e, epibranchials proper, commonly called ceratohyals, tipped
with cartilage, /, /. — 3. Glottis, or opening of trachea in the mouth ; a, base of tongue ; 5, b,
horns of hyoid bone ; c, rima glottidis, cleft or chink of the glottis ; d, a triangular vacuity ; e,
an elastic ligament ; d and e represent an epiglottis ; /, /, a papillose surface. — 4. Larynx viewed
from before (below) ; a, thyroid bone or cartilage. — 5. Larynx viewed from behind (above) ; a,
thyroid bone ; 5, b, its appendages ; c, cricoid ; d, d, arytenoids ; e, e, anterior border of thy-
roid, to which d, d are connected by two arytenoid ligaments. — 6. Larynx viewed from right
side ; a, thyroid ; b, appendage ; c, cricoid ; d, arytenoid ; /, /, cartilage attached to arytenoid ;
g, a tracheal ring. — 7. Larynx viewed from behind ; a, thyroid ; b, b, its appendages ; c, cricoid ;

d, arytenoids. — 8, 9, 10, 11, 12. Muscles of the larynx ; 1, 1 (Fig. 8), thyrohyoids ; 2, 2 (Fig. 9),
thyroarytenoids, or openers of the glottis ; 3, 3 (Fig. 10), oblique arytenoids ; 4, 4 (Fig. 11),
thyrocricoids ; 5, 5, (Figs. 11 and 12), posterior thyrocricoids. — 13. Bifurcation of trachea ; aba,
last entire tracheal ring. — 14. Last entire tracheal ring, viewed frmn below, crossed by the
pessulus. — 15. Bifurcation of trachea, and bronchi, viewed from below ; a, pessulus, the bolLbar,
or “bone of divarication” ; b, b, next succeeding tracheal half-rings. — 16. a, b, c, d, inferior
laryngeal or syringeal muscles, not well made out in this figure ; see text. I3ut the typical
oscine arrangement (acromyodian) is perceived, inasmuch as anterior («) and posterior (d) in-
trinsic muscular masses go to ends of the first tracheal half-ring, at b and c ; the extrinsic slip e
passing to sternum ; compare Fig. 1 at q. — 17. Trachea, etc., of the nightingale, nat. size.
(Compare Figs. 3, 67, 72, 73, 74.)

of the orifice, is called the semilunar membrane ; being the edge of a
partition common to both bronchi, it forms, in fact, the inner lip of
each bronchial orifice ; that is to say, the inner rima glottidis syringis,
or lip of the syringeal mouthpiece. This membrane vibrates with
the column of air, and is, in fact, one of the “ vocal chords.” Now
the bronchial rings which succeed are not annular, circumscribing
the bronchial tube, but are half-rings h, b), or arcs of circles to
be completed by membrane, which forms more or less (scarcely or
not half) of the circumference of the tube ; this membranous part,
termed the internal tympaniform membrane c to c), being on the
side of the bronchus which faces its fellow, while the hard bronchial
half-rings complete the rest of the cylinder. The membrane is
attached to the pessulus above. This accounts for the whole
bronchial tube and its vocal septum from its fellow. Now the
concavity of the upper two or three bronchial half-rings, on the
outer wall of the tube, but in its interior, is the place where is
developed a certain fold of the mucous membrane, projecting into
the tube opposite the septum, and forming the outer lip of the
syringeal glottis ; for this membranous fold, like the semilunar
membrane, is set quivering in vocalisation. The upper tracheal
rings which enter into this arrangement are enlarged and otherwise
modified. Thus are formed two “ vocal chords,” upon the vibrations
of which the harmonious or discordant notes of the bird depend.
The chords are struck by the hand of air indeed, but endless musical
variations result from the play of the muscles in increasing or
diminishing and variously combining the tension of the several
parts of the instrument. In giving four pairs of intrinsic syringeal
X

3o6

GENERAL ORNITHOLOGY

PART II

muscles (anterior external, anterior internal, intermediate, and
posterior, besides the extrinsic sternotracheales), as figured in
b, c, d, and e, Macgillivray is said to have understated the full oscine
number, which is five or six. In the raven, Owen describes five,
without counting the sternotrachealis : bronchotmchealis anticus,
anterior external ; bronchotrachealis jposticus, posterior external ;
bronchotrachealis brevis, posterior internal ; bronchialis anticus, anterior
internal ; and bronchialis posticus. The general arrangement, how-
ever, is fairly indicated by Macgillivray in where on the side of
the syrinx, the muscles are seen to diverge from the tracheal lateral
line to go to ends of the bronchial semi-rings.

The student will understand that my description is particular
only as regards the oscine syrinx; that in birds at large every
possible modification, almost, of lower tracheal and upper bronchial
rings occurs, and with various musculation, or with none. The
non-oscine rule for the muscles is, one on each side, if any ; and
insertion into mid-parts, not ends, of the bronchial half-rings. • The
latter character chiefly distinguishes the non-oscine syrinx when it
has several muscles. As to situations of the syrinx, three have
been recognised : the ordinary bronchotracheal, in formation of which
both bronchi and trachea take part; the tracheal, only known to
occur in some American Passeres, as in Thamnophilus and
Opetiorhynchus, situated wholly in the trachea, the lower part of
which is extensively membranous ; and the bronchial, wholly in the
bronchi, as in Crotophaga and Steatornis.

The Song of Birds unlocks the great secret of Genesis to those
who can hear the keynote. It is the closest approach in animate
nature to the ringing of the hydrogen bells in the physics of light.
The musical instrument figured (101, is the identical pipe the
‘‘great god Pan” first fashioned for a legacy to all time, as so
sweetly said by Mrs. Browning : —

“ He tore out a reed, the great god Pan,

From the deep cool bed of the river.

The limpid water turbidly ran.

And the broken lilies a-dying lay.

And the dragon-fly had fled away.

Ere he brought it out of the river.

“ ‘ This is the way,’ laughed the great god Pan,
(Laughed while he sate by the river !)

The only way since gods began

To make sweet music, they could succeed.’

Then dropping his mouth to a hole in the reed.
He blew in power by the river.

SEC. IV

AJVA TOMY OF BIRDS

307

“ Sweet, sweet, sweet, 0 Pan,

Piercing sweet by the river !

Blinding sweet, 0 great good Pan !

The siin on the hill forgot to die.

And the lilies revived, and the dragon-fly
Came back to dream on the river.”

But the sad sequel, felt by Keats, when poor Psyche has seen
and known, and Eros has found his wings —

“ So did he feel who pulled the boughs aside.

That we might look into a forest wide.

To catch a glimpse of Fauns and Dryades
Coming with softest rustle through the trees ;

And garlands woven of flowers wild and sweet,

Upheld on ivory wrists, or sporting feet ;

Telling us how fair trembling Syrinx fled
Arcadian Pan, with such a fearful dread.

Poor Nymph, — poor Pan, — how he did weej) to find
Naught but a lovely sighing of the wind
Along the reedy stream ! a half-heard strain
Full of sweet desolation, balmy pain.”

The blessed bluebird, “ bearing the sky upon her back,” is
burthened with the same “ light load of song ” —

Have you listened to the carol of the bluebird in the spring ?

Has her gush of molten melody been not poured forth in vain ?

Ah ! then the pulse has quickened, and a sigh, perhaps, has risen,
From the breast the bluebird’s music stirs to thoughts that lack
expression —

So tender, so tumultuous are the fancies thus aroused.

The bluebird’s song breathes gladness — breathes the sweet and
solemn triumph

Love feels when all love’s passion melts in its own fruition.
Exquisitely subtile are the chords the bluebird touches —

Chords that quiver now in ecstasy, now thrill in fond expectancy.
Now die in dreams of all that might have been.

Hers is language to interpret, and translate in accents rhythmic.

All the yearning of young love to claim his own —

Of young love that trembles on the threshold of the passions.

And shrinks before the images his ardour calls to life.

Thus to the maiden musing come thronging thoughts unbidden.
When she hears this speaking echo of the hopes that glow within;
And the tell-tale blushes redden to the rose-tint on the bosom
Of the bird that dares to breathe her secret joy.

Thus to the youth impetuous, whose life is set to music —

Let love but laugh and beckon from afar —

Fulfilment sends a greeting in the soft voluptuous languor
That steals upon the senses if the bluebird’s song be heard —

3o8

GENERAL ORNITHOLOGY

PART II

This song of wondrous gladness, ever bubbling, welling, gushing.
From a fountain full of promise, inexhaustible, divine !

Sweeter far these liquid accents when the buds of hope are blighted.
And the tree of knowledge bears its bitter fruit ;

When memory sits brooding on the ashes of her birthright.

And sackcloth shrouds a heart that once was young ;

For a silver chord is quickened where was greedy, silent sorrow —
Kesponding to a sympathetic touch :

The bird sings true and tender, with a precious burden laden.

With the tidings of a love that never dies.

So in the timid spring-time, when the world wears wreaths of roses,
King clear the joyous melodies of hope !

So in the summer season, when the wine of pleasure reddens.

Ring passionate the triumphs of the heart !

So in the sad, still autumn, when life bends beneath its burden.
When what might have been has never come to pass.

Rings once again this music on the crushed and wounded spirit,
Bringing light where all was dark and drear before :

All is not lost if the music that the bluebird bears be heeded,

For her mission is to tell us love is God.

Though it is a fact that “the Chenomor^plm are not provided
with intrinsic syringeal muscles,” there may be much truth in
treatises de canfu Cycni morituri which have appeared from time to
time, and to the number of which I may be pardoned for adding —

How sadly sweet, how soft and low
Is the music born of pain —

How mournful sounds the ebb and floM%

What measured beats, what throb and throe,

In the wild swan’s dying strain !

The archer. Death, and the twanging bow.

And the fateful shaft on-sped,

All state and grace and pride laid low.

Disordered plumes and crimson flow —

For the white swan’s heart has bled.

But hear the mournful cry that rings
On the startled air of night !

As a spirit form in the darkness wings
Its way unseen, the wild swan sings
His psalm of life and light.

How sadly sw'eet the solemn strain —

The dirge of the dying swan !

That wondrous music, child of pain.

That requiem, sounding once again —

And a bird’s soul passes on.

SEC. IV

ANATOMY OF BIRDS

309

/. Splanchnology : the Digestive System.

The Alimentary Canal, or digestive tract, is a tube which passes
through the body from mouth to anus, conveying food, the nutritious
qualities of which are drawn off by the lacteals in transitu and
assimilated, the refuse being voided. This is digestion. The canal
is really a tube within a tube, being contained in the cavity below
the bodies of the vertebrae, formed by the series of hcemal arches.
Birds are very fast livers, their digestive operations, like the pro-
cesses of respiration and circulation, being very active and effectual ;
they require proportionally greater quantities of food. The voracity
of the cormorant is proverbial, but it is probably not greater than
that of the ethereal nightingale. Birds as a class are omnivorous ;
many species are as nearly omnivorous as any animals can well be ;
but the majority are either vegetarian or flesh-feeding. Very many
birds feed upon fruits, hard or soft ; but even these, when in the
nest, are nourished for the most part upon the bodies of insects ;
and it may be truly said that the great majority of birds are
insectivorous. Birds seem to be the great controlling agency in the
economy of nature of the increase of insect life ; agriculture would
be difficult if not impracticable without them, and their economic
value is simply incalculable. Insectivorous birds cannot be much
interfered with, without destroying one of the most important and
consequential of nature’s many beautiful adjustments. The bird
cries perpetual “ 4chec ! ” to the insect. Even those birds which
are mainly flesh-eaters, as the hawks and owls, are similarly beneficial,
for the creatures they chiefly prey upon are the small rodents so
hateful to husbandry. The carrion-eaters contribute largely to make
tropical regions habitable to man. Various tribes of birds feed
almost exclusively upon fish ; and these sometimes reach the dignity
of diplomatic and other political interests of mankind : nations have
gone to war over the dung of such birds, guano-beds being to some
of the South American powers a large item of their revenue. Chili
and Peru have been fighting lately, and the United States have been
wrangling over the excrements of the alimentary canal of sea-birds.
This tube in general is shortest, simplest, and most direct in the
flesh- and fish-eaters, the nature of whose food assimilates already
more nearly to the substance of their bodies than does that of the
vegetarians. The tube is modified in different portions of its extent
for the prehension, retention, saturation, maceration, and comminu-
tion of food, and the mixture with it of other solvent fluids than
those secreted by the mucous membrane of the alimentary canal
itself. Hence arise the various modifications of its length, dilatation
here, contraction there ; the presence in its lining membrane of
numerous follicles ; and the annexation of various glandular organs.

310

GENERAL ORNITHOLOGY

PART II

Being always longer than the body, the tube is necessarily coiled
away in certain places ; this folding taking place chiefly in the
intestinal part of the tract. Modifications of structure make recog-
nisable parts, as the mouth, gullet, crop, stomach, gizzard, intestine,
cloaca, anus. Annex organs are the salivary glands, the liver, and
the pancreas, all of which pour their secretions into the canal. This
tube also receives the terminations of other systems of organs : the
auditory organ of special sense ; the respiratory system, which
is at first a mere bud or offset from the digestive; the urinary
and the generative, which, though originally distinct, primitively
and permanently open into the lower bowel. The intestine is also
continuous with the cavity of the umbilical vesicle of the embryo,
a primitive structure which disappears as the chick matures ; and
with that of the allantois, another embryotic organ which begins by
budding from the intestinal cavity. Its connection with the system
of blood-vessels is direct through the lacteals and thoracic ducts
(p. 295). Its operations are automatic and spontaneous, of the
“reflex” order; that is, excited by the presence of food, — having
work to do making it work, so to speak. Its innervation is chiefly
by the pneumogastric and sympathetic nerves ; and digestion is the
most purely vegetative function, dealing with the raw materials of
nutrition and consequently of the growth and repair of the whole
body. The active factors in this transaction are several species or
varieties of small creatures, called Entemmoehm ; they are all derived
by descent with modification from the hypoblastic cells of the early
embryo. Those of the canal itself form all the mucous epithelium
of that structure, with its various secretory crypts, follicles, and villi ;
similar creatures, perhaps of different genera, form the lining of the
salivary, hepatic, and pancreatic glands. Blood-vessels in intimate
connection with the digestive organs form that special venous
arrangement by which the blood coming from that part of the
intestinal tract where chyle is made is collected in a ported system
and sent through the liver, — in the embryo a sort of “ great dismal
swamp ” which interrupts the ordinary current. The tube within the
tube is fixed not only at its ends, but by various membranous con-
nections, among them the mesenteries. We will notice the several
departments of the alimentary canal and its annexes ; reference
should be made to Fig. 101, where most parts of the digestive system
are shown.

The Mouth and Tongue. — The most anterior of the special
cavities into which the tube is divided, and the “ manual ” organ
it contains. The mouth in general corresponds to the shape
of the jaws, already sufficiently noted. The anterior part is
much hardened, like the beak ; in fact, this hardness of the buccal
cavity, and the absence or very slight distinction of a “ soft palate,”

SEC. IV

ANATOMY OF BIRDS

311

are among the peculiarities of a bird’s mouth. There is consequently
little distinction, if any, between mouth proper and fauces, or pharynx,
which is the posterior part, leading directly into the gullet. Be-
sides this communication, the mouth receives the terminations of
four special cavities. 1. The posterior nares, on the roof of the
mouth posteriorly, generally a median slit leading into the nasal
chambers. 2. The generally single and median and more posterior
opening of the Eustachian tubes, which lead
into the tympanum, and are the remains of
the first postoral visceral cleft of the early
embryo. 3. The glottis (Fig. 101, c), a

slit at the base of the tongue, the opening
of the windpipe, and so of the whole
respiratory system, which is defended by
a rudimentary trap-door, the epiglottis, if
any. 4. One or several pairs of orifices,
the openings of the ducts of the salivary
glands. These structures, corresponding
to the parotid, submaxillary, and sublingual
glands of mammals, vary extremely in
their development. In woodpeckers, for
example, and some Raptores, elaborate
special salivary glands occur, having a
glomerate structure and a special duct of
Stenson. In many other birds, similarly
compound but less elaborate submaxillary
glands pour their secretion into the mouth
by a series of pores. In most birds, how-
ever, the salivary glands are small, simple,
and less distinct from various other sets of
mucous crypts which open into the mouth.

In the great bustard {Otis tarda; Fig. 102)

there is a singular buccal structure ; a great from Gah’od. " a, tongue ; &, the
pouch opening beneath the tongue, suscep- in front of c, the trachea, behind
tible of distension during those amatory oesophagus, a, with

antics termed the “ showing - off ” of

the creature. It is in fact an air-sac, but not of the kind already
considered (p. 296), having no connection with the respiratory
system. The narial, Eustachian, and glottidean apertures are com-
monly defended by retrorse papillae ; and other such processes of
mucous membrane, knobbed or acute, may occur elsewhere in lines
and patches. The roof of the mouth is nearly all “hard palate,”
as already said ; its soft floor is the mucous membrane and skin
between the forks of the under jaw, with muscular or other interven-
ing structures. The principal flooring muscle is the mylohyoid;

Fig. 102. — Gular pouch of
bustard ; copied by Shufeldt

312

GENERAL ORNITHOLOGY

PART II

the geniohyoid (Fig. 101, t/) is another, which passes like the first

from the mandibular to the hyoid bone ; a third is the stylohyoid
{e). The floor in some cases forms a pouch, which, as in the peli-
can, is of great extent and susceptible of enormous dilatation.

The handler of the mouth, or lingual organ, is the tongue, which
answers the same purpose as in other creatures : it is tactile, to some
extent gustatory, sometimes prehensile, nearly always manipulatory.
In some birds, as the pelican and ibis, and also the kingfisher, it is
very slightly developed, — scarcely more than a pad at the bottom
of the mouth, enjoying the most limited motion or other function.
In some birds, as the parrot and duck tribes, and also the flamingo,
the tongue is large, thick, and fleshy, quite filling the mouth. In
the first-named of these, it is dexterously manipulatory ; the morsel
of food is managed between the tongue and upper beak ; the tactile
certainly, and perhaps the gustatory sense is highly developed ; and
the fleshiness of the tongue may affect that power of articulate speech
for which some parrots are justly noted. In the Lamellirostres just
mentioned the tongue has lateral processes corresponding to the
denticulations of the beak, and the under surface is horny at the
end, like a human finger-nail. In the woodpeckers (Figs. 73, 74)
the tongue itself (glossohyal part of the hyoid) is reduced to a
slight horny and spiny tip of the lingual apparatus ; but other parts
of that mechanism are so extraordinarily developed that the “tongue”
appears as a lumbriciform (worm-like), spear-headed organ usually
capable of great protrusion from the mouth, and therefore acting as
a prehensile instrument, being bedewed for that purpose with tena-
cious saliva from the great salivary glands ; while it is actuated in
protrusion and retraction by specially developed muscles. In the
snipe and many other long slender-billed waders, the tongue is
similarly slender, but not protrusible. The long narrow tongue of the
toucans {Rhamphastidce) is beset with slender processes, so that it
seems feathery. The tongue of the humming-bird is very singular.
These and other interesting extremes aside, the ordinary style of a
bird’s tongue is flat, narrow, more or less sagittate or lanceolate,
and tipped or sheathed in horn, commonly with lateral backward
processes like the barbs of an arrow-head, — the whole glossal struc-
ture upborne pretty distinctly upon the end of the basihyal bone.
(See Fig. 101, where i, a, is such an ordinary tongue, and a-f, is its
whole skeleton.) Such horny tongues are commonly bifid at the
extreme tip or there variously lacerate, or laciniate, or thready, —
and even the fleshy tongue of some parrots, as the lories, is brushy
at the end. The bony foundation of the tongue is the composite
hyoid bone, already often mentioned ; the free lingual part proper
is based upon the glossohyal and its terminal cartilage ; the roots
curve more or less extensively about the base or more of the skull.

SEC. IV

ANATOMY OF BIRDS

313

The tongue is moved by some intrinsic muscles, as well as by those
extrinsic ones by which it is connected to the skull, jaw, and wind-
pipe (Fig. 101, ^ and ^).

The (Esophagus. — After comminution, if any, by the beak, and
insalivation in the mouth, food passes directly through the pharynx
into the oesophagus or gullet, — a musculomembranous tube connect-
ing mouth with stomach (Fig. 101, \ g, A, i). This is composed
(besides its mucous membrane) of circularly disposed constrictor fibres,
and longitudinal contractor fibres, of Myamoeha, of the pale, smooth
species {M. Icevis). It has generally a pretty straight course, but
may be diverted to one side or the other ; and, in particular, is
subject to various dilatations and contractions, permanent or tem-
porary, aside from the mere distension caused by the passage of
food. When the floor of the mouth is wide and loose, the gullet
partakes of the same character above ; the extreme case is afforded
by the pelicans, especially Pelecanus fuscus. But the gullet of many
small birds, as various genera of Fringillidce and Corvidce, is much
more distensible than is commonly supposed, and may be found
crammed with seeds which there find resting-place for some time.
The fish-eating birds, as herons, cormorants, loons, and others, have
also capacious gullets. The Australian bustard, Eupodotis australis,
has an oesophagus capable of such extraordinary distension that it
hangs down in front of the breast when inflated with air, as it is in
the amatory display in which that species is wont to indulge.
Aside from mere distensibility of transient character, the oesophagus
of man}^ birds becomes modified anatomically into a special pouch,
— the crop or craw, ingluvies, where the food is detained to be macer-
ated in a special secretion before passing on to the true stomach.
Such definite crops occur in birds of prey, which gorge such masses
of food in their irregular voracious banquets that it cannot all be
received into the stomach at once ; and likewise throughout the
orders of Columbine and Gallinaceous birds, which habitually feed
upon seeds and other fruits so hard that they are advantageously
macerated as a preliminary to true digestion. The common fowl
furnishes a good illustration of a large, definite, single and median
crop ; in pigeons it is a pair of lateral dilatations. In these latter
birds, when they are rearing their young, the secretion of the inglu-
vies, always copious, becomes still more so, and of a milky character
in consequence of the activity of the altered mucous surface ; it is
regurgitated into the mouths of the young along with the macer-
ated grains. “ This phenomenon is the nearest approach in the
class of Birds to the characteristic mammary function of a higher
class ; and the analogy of the ‘ pigeon’s milk ’ to the lacteal secre-
tion of the Mammalia has not escaped popular notice.” Various
other birds also feed their young by regurgitation of elaborated

314

GENERAL ORNITHOLOGY

PART II

food ; and very many similarly reject indigestible portions of their
ingesta. Such vomiting is best known to be the wont of birds of
prey, which habitually throw up the hair, feathers, and bones of
their victims, made up into the boluses called “ castings ” \ but the
practice is far from being confined to these flesh-eaters. The ex-
treme case of emesis offered by birds is witnessed in the hornbills
{BucerotidcB) which have been known to throw up the coat of their
stomach without discomfort, — what a blessing it would be to some
old topers if they could do the same and grow another with equal
ease ! In fact, in consequence of the capacity and directness of the
gullet, vomiting is very easy to birds, and with some it is a means
of self-defence, — very effectual, for instance, in the cases of American
vultures (Cathartides). Fish-eating birds, as herons, gulls, petrels,
habitually vomit when wounded or otherwise molested.

The Proventrieulus. — The tube just considered ends below in
a special tract, variously dilated or not, but always peculiar in the
presence of certain gastric follicles which secrete the digestive fluid
proper. The “ stomach ” of a bird, in fact, is compound, consisting
of a glandular or digestive portion, and a muscular or grinding part.
The former is the proventrieulus ; whatever its size or shape, or
whatever its magnitude in comparison with the grist-mill, it is re-
cognised by the presence in its mucous surface of these gastric
follicles, secreting the peptic fluid which chymifies the food. The
follicles are perhaps always large enough for this part of the tube
to be recognised by the naked eye, — the mucous membrane having
here a thickened, velvety, vascular appearance. The glands are of
various sizes and shapes, — usually simply tubular, sometimes clubbed
or conical, or variously racemose (like a bunch of grapes). They
are disposed in a zone around the tube, or in patches upon part of
its surface, — in the darter (Plotus), very singularly in a separate
lateral compartment looking like a crop. Details of the grouping of
these solvent glands are interminable. Whatever its anatomical
variations, and however like the end of the oesophagus it may
simply appear to be, this ventriculus glandulosus is the bird’s proper
stomach (Fig. 101, ^,j).

The Gizzard. — Mixed with the salivary, ingluvial, proventri-
cular and other secretions of the mucous surface, and already
chymified, the food of birds next passes directly into the gizzard,
gigerium, or muscular division of the stomach, sometimes called the
ventriculus bulbosus. The two are sometimes separated by a tract,
sometimes immediately consequent. In the muscular gizzard, the
food-grist is ground fine. To this end, the walls of the cavity be-
come developed into a more or less powerful muscular apparatus,
and the mucous membrane changes to a tough, thick, horny, occa-
sionally even bony, lining ; this callous cuticular lining being often

SEC. IV

ANATOMY OF BIRDS

315

very loosely attached, and even deciduous in some cases. The
muscular arrangement is chiefly in two great masses, called the
lateral muscles, converging to a central tendon ; between them inter-
mediate fibres may form a more or less distinct muscular belly. In
the most powerful gizzards, the muscular tissue is very dense and
dark coloured, the tendons brilliantly glistening, and the contained
“ millstones ” extremely callous. Such a gizzard is well displayed
by the common fowl or the goose. The opposite extreme is afforded
by the carnivorous and especially the piscivorous birds, whose soft
food requires little trituration, — it is all a matter of degree. How
readily this part of the canal responds to the regimen of the bird
is witnessed in the cock-of-the-plains (Centrocercus urophasianus) —
a bird whose gizzard is so slightly muscular as to appear like a
membranous bag, though its gallinaceous relatives have extremely
strong grinders. Its food is chiefly the buds and leaves of the wild
sage {Artemisia) and grasshoppers. Increased muscularity of the
gizzard has even been artificially produced. Birds whose grist is
heavy habitually swallow gravel, that these small stones may
mechanically aid in the grinding process. The action is so energetic,
that in auscultating a fowl when the mill is in full blast, the noise
of the grinding can be distinctly heard. The pebbles, in fact, have
a function which leaves “ hens’ teeth ” not merely mythical. The
kind of motion impressed upon the opposing pads of cuticle is
alternating, — a rubbing back and forth to a slight extent. Peculiar
dispositions of the callous surfaces are found in some pigeons, with
corresponding peculiarity of the cross-section of the gizzard. In
some of the cuckoos a matting of impacted hairs of lepidopterous
insects has been mistaken for a coat of the gizzard itself. In the
darter, which has a pyloric division or compartment of the gizzard,
this is nearly filled with a mass of matted hairs, a peculiar modi-
fication of the epithelial lining, serving to guard the pyloric orifice.
Folds of the lining membrane form a pyloric valve in many birds.
The pylorus, or the pyloric orifice, is that opening by which food
leaves the gizzard for the intestines ; the orifice of entrance from
the oesophagus is the cardiac. The two are always near together,
and sometimes adjoining. (In Fig. 101, ^ is on the central tendon

of the moderately muscular gizzard ; the cardiac orifice is between
j and h, and pylorus between I and h.)

The Intestine continues the alimentary canal to the cloaca.
Any difference in the length of the whole tract, relatively to that
of the bird, is chiefly produced by the foldings of the intestine,
especially in the upper portion of its course. The extremes of
proportionate length are perhaps not ascertained ; but known to be
from less than 2 : 1 to more than 8:1. In birds there is little or
no distinction between “ small ” and “large” intestine as to the

3i6

GENERAL ORNITHOLOGY

PART II

calibre of the tube, nor is the latter sacculated as in mammals.
The former is considered to extend from the pylorus to the cma
(structures to be presently noticed). Above the caeca the intestine
commonly receives its foldings and windings ; below them it usu-
ally proceeds more directly, or quite straight, to the cloaca, forming
literally a “rectum”; but in the ostrich this ultra-caecal tract is
longer than the rest, and convoluted. The cis-caecal portion is con-
ventionally divided into duodenum, jejunum, and ileum; there is,
however, no positive anatomical distinction of these parts in any
animal with which I am acquainted. In birds, a “ duodenum ” is
perhaps as distinct as ever ; it forms the most constant duplication
of the intestine, the pancreas being lodged in this duodenal fold (Fig.
101, \ I, m, n). The course of the intestine is otherwise very vari-
ous in different birds. The upper end, near the pylorus, receives the
hepatic and pancreatic ducts ; and food is chylijied after impregna-
tion with the biliary and pancreatic fluids ; a process furthered by
the proper secretions of the intestinal follicles. The chyle is drawn
off by the lacteals already described (p. 295), and the unassimilable
refuse of the food becomes excrementitious.

Caeca (Lat. ccecus, blind, in the nom. pi. cma; sing, caecum). —
The “ blind guts,” so called because they end in culs-de-sac, are of
two kinds. One is the umhilical caecum, or mtelline caecum, a rudi-
mentary, or rather vestigial, structure, the remains of the open duct
by which the cavity of the umbilical vesicle (an embryonic organ)
communicated with that of the intestinal tract. It is ordinarily
not to be noted at all ; but it is said by Owen to have been found
half an inch long in the gallinule, an inch in the bay ibis, and
dilated into a sac an inch in diameter in the Apteryx. The
structures ordinarily called caeca, or caeca coli, for they are usually
paired, are pouches or diverticula which set off from the intestine
proper at the junction of the ileum with the colon ; but there is
nothing in the intestine itself to mark this point, so that when
cseca are absent, as frequently happens, no distinction of ileum from
colon or rectum is appreciable. No other part of the intestinal tract
is so variable as the csecal ; so that presence or absence of these
appendages furnishes zoological characters nowadays taken very
commonly into account in framing genera and families. There are
no cseca, as in the turkey-buzzard and some pigeons ; there is a
single small caecum in herons. From a condition of extremely
small size, like little buds upon the intestine, cseca are found to
elongate to extraordinary dimensions ; and the large specimens are
frequently saccate or clubbed, with slender roots. In geese and
swans the cseca are a foot long, more or less ; in some grouse they
are said to be a yard long. In the ostrich the mucous membrane
is thrown into a spiral fold. However developed, the physiology

SEC. IV

AJVA TOMY OF BIRDS

317

of these intestinal appendages is the detention of food until all its
nutritive qualities are absorbed, and increase of the absorbent
surface.

The Cloa'ea (Fig. 101, \ or “sewer,” very well named, is the
termination of the bowel, — an oval or globular enlargement of the
rectum, of sufficient capacity at least to contain the completely
shelled egg. For, not as in placental mammals, the urogenital and
digestive organs are behindhand in their evolution, and do not
entirely lose connection with each other. Nor is there in birds
any distinct bladder ; but a cavity, originally that of the allantois
of the embryo, persists in common with that of the intestines, and
is the cloaca. Such incomplete distinction between the two as there
may be, by a folding of mucous membrane or partial compartment
of the whole, results in cloaca proper, and urogenital sinus, in which
latter are the papillose orifices of the ureters, one on each side,
from the kidneys ; and of the single oviduct ( $ ) or paired sperm-
ducts {$), from ovary or testes. The urine of birds not being
liquid requires no more of a bladder than this sinus furnishes. The
same cavity contains the penis of those birds, as the ostrich and
drake, which are provided with an organ of copulation. A peculiar
anal gland, the hursa Fahricii, also opens into the cloaca. Eefuse of
digestion, the renal excretion, the spermatic secretion, and the
product of conception, are discharged by a single anal orifice, the
two former en masse.

Being intimately related to dietetic regimen, and so to the
habits of birds, the alimentary canal varies greatly, — even more
than my slight sketch shows, — and consequently affords good
zoological characters in the details of its construction. But of all
the anatomical systems, this is the one most variable as a matter of
physiological adaptation (see p. 103). Its characters, even when they
seem weighty, are therefore peculiarly liable to be fallacious as
indices of natural affinities, and must be applied with discreet
caution to morphological classification. Such are commonly only of
generic significance. Thus in pigeons the caeca and even the gall-
bladder may be present or absent in neighbouring genera.

Alimentary Annexes. — Some of these, as the salivary glands,
have been noticed already. The two most important bodies
connected with the digestive tract, and properly considered adjuncts,
are the pancreas and the liver. The former is that kind of
lobulated salivary gland which in mammals is called the “sweet-
bread.” It lies in the duodenal loop, along which its loosely
aggregated lobes extend. Its ducts, formed by the successive union
of smaller efferent tubes, are two or three in number ; they pierce
the intestine a little below its commencement at the pylorus, and
pour into the canal the pancreatic juice, which has the property of

GENERAL ORNITHOLOGY

PART II

318

emulsionising fat. The liver is a well-known glandular organ of
very special structure and function, secreting the fluid called hile^
also received into the intestine. It is of moderate size in birds,
and deeply divided into two principal (right and left) lobes : in
some birds there is also a smaller lobe ; and one of the large lobes
may also be divided. The lobes dispart above to receive between
them the apex of the heart ; they are held in place by pleuro-
peritoneal folds contributing to form the thoracic-abdominal air-cells.
This viscus receives venous blood from the extensive portal system;
two hepatic veins then conduct it to the post-caval. The emunctory
ducts, carrying off the bile, are two or three in number. One at
least goes directly to the intestine, and another to the gall-bladder,
when that cyst exists ; in which case there is a separate cystic duct
from the bladder to the intestine, no ductus communis choledochus, or
duct common to the hepatic substance and its cyst, being formed
in birds. Two hepatic ducts may coexist with a cystic duct,
making three to the intestine, all separate ; two is the rule when
there is no gall-bladder. These emunctories commonly enter the
intestine some distance apart, and beyond the pancreatic ducts. The
gall-bladder is generally present, sometimes absent ; it may occur
or not in closely related genera of birds.

g. Oology : the Urogenital Organs

The Urinary and Generative Organs may be conveniently con-
sidered together, not only on account of their close anatomical
relations, but because their physiological functions, totally diverse
in adult life, are primitively related in the most intimate manner.
For it is a singular fact that the mean office of straining urine out
of the system is at first sustained by a structure (Wolffian body), in
closest connection with which, in the female, actually as a part of
which, in the male, are later developed those organs (ovary and
testis) whose exalted office is creative ; for these permanent genital
glands procreate the microscopic creatures called Dynamamcehce, the
marriage of which results in the reproduction of a complex
organism like the male or female parent. (See Figs. 103, 104, and
following.)

The Wolffian Bodies, or primordial kidneys, are a pair of tubular
structures which appear very early in the progress of development
of the embryo, beneath the spinal column, in front of the fore end
of the future kidneys ; with each of them is developed a duct, the
Wolffian duct, which carries their excretion into the cavity of the
allantois (the future cloaca). Upon the appearance of the true
kidneys, the transitory Wolffian bodies and ducts lose their urinary
function ; they ultimately disappear from the female, for the most

SEC. IV

ANATOMY OF BIRDS

319

part, leaving only a trace of their former existence in certain
vestigial structures {parovaria, etc.) j in the male, likewise, they
atrophy, hut not to the same extent ; for a portion of the bodies
persists as an accessory (epididymal) portion of the testicle, and
their ducts persist as the sperm-ducts, or vasa deferentia. Mean-
while, in closest connection with the Wolffian bodies, appears a pair
of organs, the genital glands, for a while exactly alike. If the new
creature is to become female, the genital gland develops to a certain
complexity of tissue and becomes the ovary ; while a certain duct,
the Mullerian duct, developed coincidently to connect such ovary
with the cloaca, becomes the oviduct. In birds usually only one
ovary and oviduct (the left) becomes functional. If the new
creature is to become mode, the same genital gland develops to a
higher degree of complexity, acquires a tubular structure, and
becomes the testicle ; it connects with remains of the Wolffian body,
and the Wolffian duct becomes the permanent sj^erm-duct, conveying
the product of the male function to the cloaca, just as the oviduct
conveys the product of the female function to the same sewerage. Thus
the testicle of the male and the ovary of the female are homologous,
in fact primitively identical organs, upon which sexual difference is
impressed by the greater complexity of structure acquired if the sex
is to be male ; a female being, anatomically and physiologically,
simply an imperfect male, arrested at one stage of her physical
progress to male perfection of structure ; and the whole nature of
the female bears out the same relation of inferiority. But the
oviduct of the female and the sperm-duct of the male, though
physiologically identical, having the same function of conveying
the products of generation from the genital gland to the light of
day, are not anatomically the same ; for in the case of the female,
whose Wolffian duct has disappeared, the Mullerian is the oviduct; in
the case of the male, in which no Mlillerian duct appears, the
Wolffian is the sperm-duct. The two are analogous, not homologous
(a good illustration — seepage 103). But it must be further observed
that while the sperm-duct conveys only the masculine essence from
centre to periphery, the oviduct conveys the feminine material from
centre to periphery, and also the male essence in the opposite
direction ; for upon coitus, which is direct in all birds, the sperma-
tozoa deposited in the cloaca of the female find their w^ay up
through her oviduct to the ovary, there to accomplish impregnation
of the ovarian ova, the fecund product then passing down by the
same avenue. All that relates to the mysteries of generation — both
the structure and function of the reproductive organs, and the
maturation of the product of conception, is properly Oology (Gr. wov,
oon, an egg) ; though the term is vulgarly used to signify merely a
description of the chalky substance in which the egg of a bird is

320

GENEJ^AL ORNITHOLOGY

PART II

finally invested. The anatomy of the egg is Embryology. An egg,
or ovum, is simply the product of conception up to the time that
product acquires an independent existence ; while still connected
with the female tissue of the ovary, and before or after it amalgam-
ates with the male element, it is an ovarian ovum; more or less
incompletely matured, it is an embryo or foetus — the former term
being commonly applied to the unhatched young of birds. The
only difference between the “ egg ” of a “ viviparous ” mammal and

Fig. 103. — Urogenital Fig. 104. — Urogenital organs of Fig. 105. — Urogenital
organs of male embryo bird ; female embryo bird ; from Owen, organs of tlie domestic
from Owen, after Miiller. a, after Miiller. a, kidneys ; Wolf- cock; after Owen, a,
kidneys ; b, ureters ; c, Wolf- fian bodies; c, genital gland, to testis; b, epididymis ; c,
fian bodies ; d, their ducts, become ovary ; d, adrenals ; e, ure- sperm - duct or vas de-
to become sperm-ducts ; e, ters ; /, Wolffian ducts, to disap- ferens ; d, adrenal ; k,
genital glands, to become pear ; g, Mullerian ducts, to become cloaca ; x, kidney ; y,
testicles ; /, adrenals. oviducts. ureter.

that of an “ oviparous ” bird is in the albuminous and cretaceous
envelopes of the latter, and its speedy expulsion from the body of
the female to be hatched outside, without anatomical connection
with the mother after the hard shell is formed ; whereas in most
mammals the ovum is retained in a dilated part of the Mullerian
duct (uterus or womb) until it “ hatches ” ; but mammal and bird
alike “ lay eggs,” the essential germinative part of which is identical.
Appreciation of these facts, and a proper idea of the relations of
the mature sexual organs to the Wolffian bodies, is necessary to
any understanding of the parts and processes concerned in repro-

SEC. IV

ANATOMY OF BIRDS

321

duction.^ We have here to consider the permanent as distinguished
from the transitory kidneys, and may then recur to the subject of
generation.

The Kidneys (Lat. renes, Engl, reins^ adj. renal ; Figs. 103, 104,
a ; 105, x) differ much from those of mammals in physical
characters, though identical in function — that of straining off from
the blood certain deleterious substances in the form of urea ;
whence they are sometimes called emulgent organs. Their office of
purification is analogous to that of the lungs, which decarbonise the
blood, and to some extent vicarious, as is that of excretory organs
in general. As the lungs are closely hound down to the thoracic
region of the trunk, so are the kidneys impacted in the pelvic
region, being moulded to the many sacral inequalities of surface.
They are paired, but sometimes connected across the median line
by renal tissue ; they have no special renal artery, but derive their
blood from various sources ; and blood from them takes part in the
hepatic portal system, no reniportal being accomplished. They have
little or nothing of the particular mammalian configuration which
has made “ kidney-shaped ” a common descriptive term ; being
elongated, somewhat parallel-sided, and rectangular, flattened bodies,
lobated into a few large compartments, and lobulated into many
lesser divisions ; their figure depends much upon that of the pelvis.
They are very dark coloured, rather soft, easily lacerable, and
appear to the naked eye to be of a granular substance, without
distinction of “cortical” and “medullary” portions. Nor is there
any “ pelvis ” of the kidneys in which the uriniferous tubules empty
together by numerous ducts as in a common basin. Each ureter
(Figs. 103, h; 104, e; 105, y) or excretory duct is formed by
reiterated reunion of the tuhuli uriniferi, after the manner of a
pancreatic duct; each ureter passes down behind the rectum and
opens into the lower back part of the cloaca — much like a
mammalian ureter into the base of the bladder. The original
cavity of the allantois remains to furnish no more of a urinary
Uadder than some special dilatation of the cloaca represents ; but
this rudimentary bladder, as distinguished from the urogenital
sinus in which the ureters terminate alongside the sperm-ducts, is
well marked in some birds ; being in the ostrich, for example, a
considerable enlargement of the cloaca between the termination of

^ The matter may be further illustrated by the two figures borrowed from Owen
(after Muller). In both figures, the large dark masses, a, are the permanent kidneys,
whose ducts, h in Fig. 103, e in Fig. 104, are the ureters, emptying into the clo^a. In
Fig. 103, male, c is the Wolffian body, whose duct, d, persists as the sperm-duct, con-
veying semen from e, the testis. In Fig. 104, h is the Wolffian body, whose duct,/, dis-
appears ; and g is the Mullerian duct, becoming the oviduct, to convey the egg from c,
the ovary. Thus e, Fig. 103, and c, Fig. 104, are the homologous genital glands, becoming
either testis or ovary ; but the sperm-duct, d, Fig. 103, is not the oviduct, 5^, Fig. 104.

Y

322

• GENERAL ORNITHOLOGY

PART II

the rectum proper and the urogenital compartment of the sewer.
The renal excretion is not watery as in mammals, but semi-solid,
and voided with the faeces, of which it forms part.

The kidneys are capped by a pair of small yellowish bodies, the
sujprarenal capsules or adrenals (Figs. 103, f; 104, 105, d), the
nature of which is undetermined. They are chiefly interesting to
the practical ornithologist from their liability to be mistaken for
testes in examining specimens for sex (see p. 68).

Male Organs of Generation. — The testis (Lat. testis, pi. testes,
a witness ; Fig. 105, a) or testicle has been already sufiiciently noticed
as to its general appearance and position (p. 69). As said above,
it is the essential male organ, consisting of the primitive indifferent
genital gland (Fig. 103, e) in its highest state of development as a
tubular secretory organ, connected with the remains of the Wolfiian
body as a part of its efferent structure (epididymis; Fig. 105, h) and
with the original Wolffian duct as its ms deferens (Figs. 103, d, 105,
c), or efferent duct, by which the semen is conveyed to the cloaca.
The original glands normally remain paired, and both are usually
functionally developed to corresponding size, shape, and activity ;
they remain in their embryonic situation in front of the upper part
of the kidneys ; and such difference of appearance as they present
under different circumstances is mainly seasonal. For birds, as a
rule, procreate only at particular times of the year, rarely having
more than one or two broods of young : the functional activity and
quiescence of the testes correspond, as the enormous swelling of
the gland during the breeding season is olie of the peculiarities of
the bird’s organ. This may be related to the absence, in birds, of
specially formed vesiculoe seminales, or seminal reservoirs ; though
certain contortions and dilatations of the sperm-ducts which are to
be observed may imperfectly answer to detain the secretion until
circumstances render it available. The passage of the sperm-duct
is along the face of the kidneys, generally in company with the
ureters ; the opening is by a papilla upon the surface of the uro-
genital sinus. These papillose terminations of the sperm-ducts
are erectile to a degree, and answer the purpose of paired penes in
those birds which are not provided with better-formed copulatory
parts. In coitu the cloacal chambers containing the orifices of the
genital ducts are opened, and the more or less protruded papillse
come in contact or close juxtaposition. In cases in which a penis
or two penes are developed, the urethral passage is a groove, never
a tube, though cavernous and even muscular tissue may be de-
veloped ; and in any case of such an intromittent apparatus, it has
cloacal invagination when not operative. These organs, in all their
variety, are of the sauropsidan, not mammalian, type ; though in
some respects the structure approaches that seen in the non-

SEC. IV

ANATOMY OF BIRDS

323

placental mammals. No prostate or Cowperian glands exist in
birds.

The sole office of the testis, or .odjphoi'on masculinum, is the
secretion of semen, associate structures being simply accessory for
the conveyance of that vital substance and its transference to the
opposite sex. The seminal fluid itself is
merely the vehicle of transport of the sperma-
tozoa, in which their activity may be freely
exercised in their intuitive struggles to gain
access to their mates in the ovary. It is
literally a “ sea of life ” in which the minute
creatures swim in shoals to their destiny —
and their fate in any case is death. If they
successfully bulfet the waves of fate they find
a watery grave in the ovum at last; if that „(
haven be not reached they simply perish in magnified ; from owen, after
mid -ocean. I he spermatozoa, or seminal

animalcules, or male Dynamamcebm (Figs. 106, 107), are the
exact counterparts of ovarian ova, in so far as they are single-celled
animals of a very low grade of organisation ; but their activity and
intelligence are marvellous, and still more so is the mysterious attri-
bute with which they are endowed of assimilating their protoplasmic
substance with that of the ovum ; with the result that the thus
fecundated ovum is capable of procreating itself
by fission for a period until a mass of similar
creatures is engendered ; from which mass is then
speedily evolved the complex body of the Bird.
The corresponding female Dynamamoelce (ovarian
ova) are simj^le spherical animalcules, physically
indistinguishable from an ordinary encysted Amcela;
but the spermatozoa are remarkably distinguished
in appearance, furnishing probably the best marked
matozoa of ^s^?row’ case of sexual characters to be found among the
P">'otozoa, to whicli class of aiiimals they belong. The
Wagner and Leuck- spermatozoa resemble flagellate infusoria or ciliated
endothelium cells, though they each have but a
single whip. They are of extremely minute size, much smaller than
their females, and filamentous ; more or less thickened and some-
times wavy at their nucleated heads, whence protrudes an excessively
delicate thready tail, endowed with great vibratory energy. They
may be likened to diminutive attenuated tadpoles, which swim by
lashing the tail in the seminal fluid. Under the microscope shoals
of these curious creatures may be seen swimming in the sea, nosing
about in search of the ovum, butting their heads in wrong places,
backing out and trying again in another direction ; with such success

324

GENERAL ORNITHOLOGY

PART II

that out of myriads a score or so may gain their end. It will be
seen that they have a long journey to accomplish ; for, liberated in
the cloaca of the female, they have to swim through the whole
length of the oviduct to the ovary. Besides such physical differ-
ence between the male and female Dynamamcehoe. as I have indicated,
they differ in their place and mode of birth ; and in this difference
lies the very gist of sex. The original indifferent genital gland
above described, arrested, as said, at a certain stage of development
and therefore female — the ovary — produces its eggs from its surface-
cells, which subside into the ovarian tissue, and are quietly packed
away there as ovarian ova, ready to ripen and awaken to impreg-
nation in due course. The same gland, further developed into a
testis, gives active birth to the spermatozoa in the tubules of its
complicated interior tissue. In the former case, the superficial cells
slowly ovulate ; in the latter, the cells lining the interior speedily
spermate; in a word, the testis is as literally viviparous as is the
ovary oviparous — and these conditions are certainly no insignificant
indices of relative development in the scale of being. The spermatozoa
appear in some animals to be set free in myriads from the walls of
the seminal tubules whence they directly issue ; in birds, they are
described as appearing coiled or otherwise packed in delicate sperm-
cells, which speedily rupture and discharge the creatures in the
current of the seminal fluid, where they take up the course and
display the energetic actions above noted. Either case has its
parallel among ordinary Protozoans ; the former corresponding to
the process of budding or gemmation, the latter to that of interior
fission and discharge of numerous progeny by rupture of the en-
velope. The final conjugation of spermatic filaments with ovarian ova
is simple fusion, such as any ordinary sexless amoeboid animal may
practise to blend its protoplasmic substance with that of another.
But there is this difference, that in the case of Dynamamoeba it is a
true sexual congress, usually polyandrous^ and still more of a one-
sided affair in that the female Dynamamoeba is at the time in a more
or less quiescent encysted state.

Female Organs of Generation. — The connection between the
male and female organs of generation is naturally so close that in
what has preceded it has been scarcely possible to speak of the
former without reference to the female counterparts. I have thus
far endeavoured to state clearly the nature of the originally sexless
genital gland ; the difference in the same gland when afterward
sexed male or female j and the character of the spermatic offspring
of the male gland. In reading that lesson the novice in such
Eleusinian mysteries must not mistake the language I have used to
describe the male Dynamamoeba, or spermatozoon, as applicable to any-
thing in the development of the female Dynamamoeba, or ovum, into

SEC. IV

AMJ TO MY OF BIRDS

325

the chick j for all said thus far only relates to the bringing of the
spermatozoon into contact with the ovum, preliminary to the initial
step of the ovum in its course of development. It is this female
Dynamamoeba — this primitive ovarian ovum, the germ of the chick,
which corresponds to and is the counterpart of the male Dynam-
amoeba, on meeting and mingling with
which fecundation is accomplished ;
the impregnated ovum being then em-
powered to take up its marvellous
march. Conjugation of the opposite
DynamamcebcB occurs either in the
ovary or upper part of the oviduct —
most probably the former. One or
several spermatozoa — usually more
than one — accomplishing their journey
up the oviduct, and finding their
affinity, insinuate themselves into the
substance of the ovum, and die there,
dissolved in amorous pain ; that is to
say, they melt into the substance of
the ovum. The now fertile result,
consisting of the mingled protoplasm
of the opposite amoebas, is to all
appearance precisely the same as the
original infecund ovum — yet there is
all the difference in the world, as the
result shows.

The general character of the ovary

of a bird has been already indicated carus. a, h, c, d, mass of ovarian ova,
/ /I rv \ • • 1 p . , in all stages of development ; 6, a ripe

(p. 69). The principal superhcial one ; c, its stigma, where the ovisac or

rliflPprPnpp in nnnpnranpp whpn fhp calyx ruptures ; (Z, a ruptured empty

Qinerence in appearance wnen hne calyx, to he absorbed ; e, infundibulum,

ovary is in functional activity from or funnel-shaped orifice of the oviduct ;

1 /) ’^cxt portion of oviduct ; g, follicular

the corresponding organ of a mammal, part of oviduct ; m, mesometry, mem-
. 111 1 • brane steadying the oviduct the re-

IS that the ova develop to such a size, ference-line, m, crosses the constricted

in ripening in the ovary before leaving o? Ihf

it for the oviduct, that the organ looks shell-forming or uterine part of oviduct,

like a bunch of grapes very large or vaginal part of oviduct, opening into

and conspicuous. The oviduct is the ''''''' '''

musculomembranous tube (modified

Mullerian duct) which conveys the ripened ovum, and in its passage
provides it with a quantity of white albumen, and finally a chalk shell.
A bird’s oviduct is the strict morphological homologue (p. 103) of a
mammal’s Fallopian tube, uterus and vagina — more accurately, of
one Fallopian tube, one-half of a uterus, and one-half of a vagina ;
for the uterus and vagina of a mammal result from the union of

Fig. 108. — Female organs of domestic
fowl, in activity; from Owen, after

326

GENERAL ORNITHOLOGY

PART II

both Mullerian ducts ; whereas in a bird only one — the left usually
— is normally developed. Functionally, the oviduct is also analo-
gous (p. 103) to the mammalian uterus, inasmuch as it transmits the
product of conception, and detains it for a while, in the initial
stage of its germination, as we shall see in the sequel ; though all
but the very first steps in the development of the chick are taken
during incubation, the egg having so hastily left its uterine matrix.
These structures — ovary and oviduct. Fig. 108 — are most con-
venientlj^ described as we trace the course of the ovum from its
origination to its maturity. This record differs considerably from
the corresponding course of events in a mammal, inasmuch as the
ovum of a bird, though primitively identical with that of any other
animal, acquires special albuminous and cretaceous envelopes which
the mammalian ovum, developed in the body of the parent, does
not require. The process is termed ovulation. Ovulation, which is
the formation of an egg in the bird, must not be confounded with
germination, which is the formation of a bird in the egg. The
former can be accomplished by the virgin bird, which may lay eggs
scarcely differing in appearance from those which have been fecun-
dated, but germination in which is of course impossible. The course
of ovulation, and afterward of germination, is now to be traced.

Ovulation. — The ovum begins as a microscopic point in the
ovary, the stroma or tissue of which is packed with these incipient
eggs. It is primitively just like any other female Dynamamoeba,
from that of a sponge up to that of a woman — a naked simple cell,
capable of exhibiting active amoeboid movements. It consists of a
finely granular protoplasm, the vitellus, or yelk, enclosed in a delicate
structureless cell- wall, the vitelline membrane, called the zona pellucidM
from its appearance under the microscope. Imbedded in the vitellus
is a nucleus, or kernel, the germinal vesicle ; in this is a nucleolus, or
inner kernel, the germinal spot. The ovum occupies a tiny space in
the ovary, the cellular walls of which constitute an ovisac, or Graafian
follicle. Now if such an ovum as this were mammalian, it would,
without material change, burst the ovisac, be received into the Fallo-
pian tube and conveyed to the uterus ; where, supposing it already
fertilised, the whole of its contents would develop into the body of
the embryo. It would therefore be holoblastic (Gr. 6Xo<s, liolos, the
whole; /lAao-rtKos, blastikos, germinative). It is different with a
bird or other “ oviparous ” animal, the egg of which has to hatch
outside the body ; for provision must be made for the nourishment
of the developing chick, thus separated from the tissues of its
mother. Such provision is made by the accumulation about the
ovum of a great quantity of granular protoplasmic substance, which
forms nearly all the large yellow ball called in ordinary language
“ the yelk ” of an egg. None of this adventitious substance goes to

SEC. IV

ANA TO MY OF BIRDS

327

form the embryo ; it is what the embryo feeds on during its forma-
tion. A bird’s egg is therefore meroblastic (Gr. meros, a part,

and jSXaa-TLKos), and we must carefully discriminate between the
great mass of yellow food-yelk, as it may be called, and a small
quantity of “white yelk,” the true germ-yelk, which alone is trans-
formed into the body of the chick. The latter forms the cicatricle,
vulgarly called the “ tread ” ; that small disc, visible in most birds’
eggs to the naked eye, which appears upon the surface of the great
yellow ball, floating in a pale thin yelk which penetrates the denser
and yellower food-yelk by a cord of its own substance leading to a
central cavity, the false-yelk cavity, around which the food-yelk is
deposited in a series of concentric layers like a set of onion-skins.
The whole mass is surrounded by a delicate structureless yelk-skin,
called the vitelline membrane (whether this
be the original vitelline membrane of the
Dynamamoeba or not; i.e. whether the
food-yelk has accumulated inside or out-
side the original zona jyellucida). All this
enormous accumulation, effecting what is
called a metovum or after -egg to dis-
tinguish it from th.Q ]grotovum, or primi-
tive state of the egg, goes on in the
ovary, and in the ovisac of each ovum ;
with the ripening of the ovum, the ovisacs
become distended to a corresponding
size, and the whole ovary acquires the
familiar bunch - of - grapes appearance.

, , , • i* i •. i 1 yelk-skin, enclosing the yellow food-

VV ith such maturation of the fruit, the yelk, which is deposited in concen-

connection with the rest of the ovary ft’

lengthens into a stalk,
which the ripe ovum
stock, like any fruit upon its stem,

ready to burst its skin and fall into the open mouth of the oviduct.
Such rupture of the Graafian follicle (ovisac), in its now distended
state known as the capsule or calyx, occurs along a line where the
numerous blood-vessels which ramify upon its surface appear to be
wanting, called the stigma : this is rent ; the ovum slips out of its
calyx, like the substance of a grape pinched out of its skin, and falls
into the oviduct. After this discharge, the empty calyx collapses,
shrivels, and ultimately disappears by absorption. (See explanation
of Fig. 108.)

The ovum thus acquires the full size of its yelk in the ovary —
becoming, as in the case of the hen, a yellow sphere an inch in
diameter.^ Notwithstanding its enormous distension with food-
1 How great this is can only he appreciated hy comparison. The human egg, on

Meroblastic ovum
(yelk) of domestic fowl, nat. size, in
section ; after Haeckel, a, the thin

or 'pedicel bv ^ cord of white yelk (here

’ N represented in black) to the central
hangs to its cavity, d'.

328

GENERAL ORNITHOLOGY

PART II

yelk, it is still morphologically a simple cell, affording the maximum
dimension of any known protozoan or single-celled animal. Enter-
ing the oviduct, the germ-yelk part of the whole mass is fertilised
by spermatozoa, unless this process has before occurred in the ovary,
and in its passage through that tube the yelk-ball becomes invested
successively with the mass of transparent albumen known as the
“ white ” of the egg, and finally by the chalk shell — both secreted
by the mucous membrane lining the oviduct.

During its functional activity, the left oviduct (there being
usually only this one) becomes highly developed, both as to its
muscular walls, which by their contractility embrace the ovum
closely and squeeze it along, and as to its mucous secretory surface.
It is supported by peritoneal folds forming a mesometry, like the
mesentery of the intestines ; its whole structure and office are quite
like those of a length of intestine. The upper end of the singularly
serpentine oviduct is dilated into an infundibulum, or funnel-like
mouth, corresponding to the fimbriated extremity of the mammalian
Fallopian tube, and constituting a morsus diaholi, or “ devil’s grip,”
which gets hold of the ovum to drag it down to the common lot of
mortals from its high ovarian birth. The infundibulum receives
from the mesentery a delicate tunic of unstriped muscular fibres,
which are so disposed as to dilate that orifice for the reception of
the ovum ; and during the venereal orgasm the mouth of the tube
is supposed to seize upon the ripest egg. The actual anatomy of
the arrangement, and the whole operation, is strangely suggestive
of one of the oldest myths respecting the serpent which bore the
egg of the world in its jaws. The mucous lining of the oviduct
consists of a layer of ciliated epithelium ; the membrane has a
different character in successive portions of its extent. Above,
when the tube is not distended with its burthen, the lining is
thrown into lengthwise folds, which lower down become spirally
disposed, and then longitudinal again before they cease. This
rugous portion of the tube is beset with mucous follicles, which
secrete “the white.” The oviduct, after contracting at a point
called the isthmus, enlarges to a calibre sufficient to accommodate
the egg in its shell ; for this is the shell-forming part, homologous
with the mammalian uterus (a sinister semi-uterus at least), lined
with large villi, and beset with the follicles whose secretions calcify

escaping from the Graafian follicle, is said to be from to of an inch in
diameter. Taking it at there would be 40,000 in a square inch, and in a cubic
inch 8,000,000. The largest bird’s egg known, that of the Mpyornis^ is said to have
a content of about a gross of hen’s eggs — 144. SupiDosing the yelk of the ^pyornis
egg to bear the usual proportion to the other contents of the shell, and allowing for
the difference in bulk between a sphere and a cube of equal diameters, there would
still be somewhere about a billion human eggs in one JEpyornis egg-yelk — roundly, a
mass of them equal to that of the germs of more than one-half of the present popula-
tion of the globe.

SEC. IV

ANATOMY OF BIRDS

329

the egg-shell, and decorate it with pigment. The rest of the tube
is vaginal, being merely the passage-way by which the perfected
ovum is discharged into the cloaca, to be expelled per anum. The
muscular walls of the oviduct consist of both circular and longi-
tudinal unstriped fibres, like those of intestine — the latter especially
in upper portions and at the infundibulum, the former more con-
spicuously below, where they form a sort of os tincce at the bottom
of the calcific portion, and a kind of sphincter mgince at the end of
the tube. A recognisable clitoris is developed in many birds.

The deposition of the white and of the shell remains to be
noticed. The first deposit upon the yelk-ball consists of a layer of
dense and somewhat tenacious albumen, called the chalaziferous
membrane (Gr. ydAafa,
chalaza; a tubercle, and
Lat. fero, I bear). As the
egg is urged along by the
peristaltic action of the
tube, it acquires a rotation
about the axis of the
tube ; the successive layers
of soft albumen it re-
ceives are deposited some-
what spirally; and the
chalaziferous membrane
is drawn out into threads
at opposite poles of the
egg. These threads, which
become twisted in opposite
directions during the ro-
tation of the egg, are
called chalazce ; they are
the “strings,” rather un-
pleasantly evident in a soft-boiled egg, which serve the im-
portant ofiice of mooring and steadying the yelk in the sea
of white by adhesions eventually contracted with the membrane
which immediately lines the shell. They are also entrusted
with the duty of ballasting, or keeping the yelk right side up. ’ For
there is a “ right side ” to the yelk-ball, being that on which floats
the cicatricle, or “tread.” This side is also the lightest, the white
yelk being less dense than the yellow ; and the chalazse are attached
a little below the central axis. The result is, that if a fresh egg be
slowly rotated on its long axis, the tread will rise by turning of the
yelk-ball in the opposite direction, till, held by the twisting of the
chalazse, it can go no farther ; when, the rotation being continued,
the tread is carried under and up again on the other side, resuming

Fig. 110. — Hen’s egg, nat. size, in section ; from Owen,
after A. Thompson. A, cicatricle or “tread,” with its
nucleus, of white germ-yelk, floating on surface of pale
thin nutritive yelk, leading to central yelk-cavity, x ; o,
the yellow yelk-hall, deposited in the successive layers,
forming a set of Jialones, and enveloped in the chalaziferous
membrane which is spun out at opposite poles into the
twisted strings, chalazse, c, c; b, V, successive invest-
ments of softer white albumen ; d, membrana putaminis,
the “ soft shell ” or egg-pod, between layers of which at
the great end of the egg is the air space, /; e, the shell.

330

GENERAL ORNITHOLOGY

PART II

its superior position as before. After all the spiral layers of soft
white are laid on, a final covering of dense albumen is deposited at
the isthmic part of the oviduct. This forms a tough tunic called
the membrana putaminis (Lat. pufamen, a peel, rind), or “ egg-pod ” ;
it is the final envelope of such a “ soft-shelled egg ” as a hen drops
when deprived of the lime required to enable her to secrete a hard
shell. In the uterine dilatation of the oviduct a thick white fluid
charged with earthy matter is exuded ; this condenses upon the
egg-pod and forms the shell. The composition of this earth is
chiefly carbonate of lime (common chalk), with some carbonate of
magnesia, and phosphates of both of these bases — thus like that of
bone as to ingredients, but in very different proportions. The shell
does not simply overlie the pod in a distinct sheet, but is intimately
coherent, the microscopic crystals or other particles of the earthy
matter being deposited in the matted fibrous texture of the pod.
The connection is most intimate in fresh eggs ; after a while, layers
of the pod separate at the butt of the egg, forming the large
air-space which every one has noticed in that situation. The
shell being very porous, readily admits air. The air-space enlarges
during incubation, and the pod becomes more and more distinct
from the shell, which latter also increases in porosity and fragility
toward full term.” The rough or smooth appearance of an egg-
shell, the pores which may be visible to the naked eye, and other
physical characters, are due to the impression made upon it by the
lining membrane of the “ uterus.” The superficial deposit of chalk
is so heavy, in some cases, as those of cormorants, etc., that it may
be scraped off without interfering with the texturally firm shell-
substance underlying. All the coloration of egg-shells, which
frequently makes them pretty objects, is simply the deposit of pig-
ment granules in or upon the shell. Such deposit may be perfectly
uniform, as it is in the bluish -green egg of a robin (Turdus migm-
torius), for instance, but it is oftener spotty — either upon a white or
a whole-coloured ground. The browns and neutral tints are the
usual colours, particularly a bright reddish-brown ; the same, lying
in instead of upon the shell, gives the grays, “ lilacs,” and “ laven-
ders ” so well known. In ptarmigan, the pigment is so heavily
deposited that the egg comes out pasty on the surface ; a sign of
“ fresh paint ! ” one must not disregard if he would not spoil the
decoration.

Oviposition. — The energy and rapidity with which the processes
involved in the manufacture of so complex a product as a bird’s egg
is now seen to be are extraordinary. A domestic fowl may lay an
egg every day for an indefinite period. It is difficult to say how
quickly an egg may ripen in the ovary ; for, during the activity of
that organ, several or many are to be found in all stages of im-

SEC. IV

ANATOMY OF BIRDS

331

maturity, and the date of the initial impulse cannot well be deter-
mined. As there is probably but one egg at a time in the oviduct,
the whole process of finishing off the yelk-ball with its chalaziform,
soft albuminous, putaminous, and calcareous envelopes may go on
in twenty-four hours, most of which time is consumed in the shell-
formation. The number of eggs matured by the human female is
or should be thirteen annually ; this is no large number for many
of the gallinaceous and anatine birds to deposit in about as many
days. But a probable average number is five or six. Defeat of
the procreative instinct from any accident is commonly a stimula-
tion to renewed endeavours to reproduce ; and very many birds
rear two or three broods annually, though one clutch of eggs is the
rule. Many, such as auks, petrels, and penguins, lay a single egg.
Two eggs is the rule in humming-birds and pigeons. Three is
normal to gulls and terns, though these often have but two. Four
is the rule among the small waders of the limicoline groups. Some
of the small Oscines lay over the average, having eight or ten ; among
these, the European sparrow, Passer domesticus, is probably the most
prolific. The parasitic cuckoos are said to lay the relatively smallest
eggs ; that of the Apteryx is said to be the largest, weighing one-
fourth as much as the bird. The usual shape of an egg has given us
the common names oval, ovate, and ovoidal, for the well-known figure.
Some, as those of owls, woodpeckers, kingfishers, and others, more
or less nearly approach a spherical shape. Eggs of grebes, herons,
totipalmate birds and various others are rather elliptical, or equal-
ended, and narrow in proportion to their length. Eggs of the
limicoline group are generally pyriform, — very broad at one end
and narrow at the other. But the eggs of all birds vary more
in size and shape than some of the devotees of theoretical oology
admit in their practice. The variation so well known in any breed
of domestic fowl is scarcely above a normal rate. The short
diameter, corresponding to the calibre of the oviduct, is less variable
than the long axis ; for when the quantity of food-yelk and white,
upon which the difference in bulk depends, varies with the vigour
of the individual, the scantiness or redundancy is expressed by the
shortening or lengthening of the whole mass. The egg traverses
the passage small end foremost, like a round wedge, with obvious
reference to ease of parturition by more gradual dilatation of the outlet.

Germination. — Leaving now all the accessory parts of an egg,
let us confine attention to the germ-yelk, or “ tread,” which is alone
concerned in the germinative process. Eecurring to the female
Dynanwbmoeha, consisting of granular protoplasm (vitellus) included
in its cell-wall (vitelline membrane) and including its nucleus and
nucleolus (germinal vesicle and germinal spot), we will trace it up
to the time it begins to take shape as an embryo chick. At first,

332

GENERAL ORNITHOLOGY

PART II

as I have observed before, it is like any other amoeba ; the first step
of development is probably a retrograde one ; for if there ensues,
when the spermatozoa melt into the ovum, the result affirmed for
mammalian ova, the original germinal vesicle and germinal spot
disappear, and the whole content of the ovum proper is simply a
homogeneous mass of granular protoplasm. In this retrograde step,
the organism, at the lowest possible round of the ladder of evolu-
tion, is called a monerula. The germinal vesicle and spot, however,
are speedily reconstructed, and the ovum looks precisely as it did
before. But observe that the actual difference is enormous; for it

Fig. 111. — Segmentation of the vitellus by discoiclal cleavage, diagrammatic, x about 10 times,
after Haeckel. Only the “tread,” cicatricle, or germ-yelk (Figs. 109, &, 110, A) is represented,
as no other part of the whole yelk-ball undergoes the process. A, separation into 2 ; B, into 4 ;
C, into 16, by 8 radial and 1 concentric furrow ; D, into many parts, by 16 radial and about 4
concentric furrows ; E, 64 radial and about 6 concentric furrows ; F, the whole tread broken up
into a mulberry-mass (morula) of cells.

now consists of the blended substance of the original ovum and of
the spermatozoa ; and in this duplex or bisexed state, before any
further step is taken, the creature is called a cytula, — the parent
cell of the entire future organism. In the former state it could
reproduce nothing, not even itself ; for it is the strange physiological
law of a Dynamamoeha that it cannot reproduce like an ordinary
cell, but must evolve an entire organism, like both of those two whose
vital forces it concentrates, summarises, and embodies, — or nothing.

The first change in the parent-cell is that by which it becomes
broken up into a mass of cells, each of which is just like itself.
This process is called segmentation of the vitellus ; each one of the
numerous resulting cells is called a cleavage-cell. The nucleus of

SEC, IV

ANATOMY OF BIRDS

333

the parent-cell divides into two ; each attracts its half of the yelk ;
the halves furrow apart and there are now two cleavage -cells in
place of the one parent-cell. A furrow at right angles to the first,
and redivision of the nuclei, results in four cleavage-cells. Eadiat-
ing furrows intermediate to the first two bisect the four cells, and
would render eight cells, were not these simultaneously doubled by
a circular furrow which cleaves each, with the result of sixteen
cleavage -cells. So the subdivision goes on until the parent -cell
becomes a mass of cells. This particular kind of cleavage, by radiat-
ing and concentric furrowing, is called discoidal^ and the resulting
heap of little cells assumes the figure of a thin, flat, circular disc.
Segmentation of the vitellus, in whatever manner it may go on,
results in a mulberry-like mass of cleavage-cells ; and the original
cytula has become what is called a morula. This process and result
are clearly shown in Fig. Ill, A-F.

The morula or mulberry-massed germ of which the “ tread ” of
a bird’s egg at this moment consists increases by multiplication of
cells, and the disc is lifted a little away from the mass of yellow
food-yelk upon which it rests, like a watch-crystal from the face of
a watch. This disposition of the greatly multiplied cells in a layer
and their coherence forms of course a membrane, — the blastodermic
membrane, or blastoderm (Fig. 112, B, b). The cavity between the
blastoderm and the mass of food-yelk is called the cleavage cavity, s.
At the stage when the blastodermic membrane and cleavage-cavity
are formed, the germ is called a blastula, or germ-vesicle,'^ and the
process by which the morula becomes a blastula is called blastula-
tion. Next, from the thickened rim, w, of the watch-crystal-like
blastula a layer of large endoderm cells (Fig. 112, C, i) separates, and
grows toward the centre : when it gets there, of course the original
cleavage-cavity, s, is shut off from the surface of the food-yelk ; a
second crystal having grown under the first one. The second
adheres to the first, obliterating the original cleavage-cavity ; the
germ is now obviously two-layered ; the rising of the inner layer to
meet the outer results in a cavity between itself and the food-yelk,
D, d. This cavity exactly resembles the original cleavage-cavity,
but it is a very different thing, being the primitive intestinal cavity.
The blastula, or germ-vesicle, has become converted into a gastrula
by the invaginating process just described, known as gastrulation.
The gastrula of a bird has the circular discoidal form which causes
it to be termed a discogastrula. This process of forming a single
blastodermic layer, with a cleavage-cavity (blastula, or true germ-
vesicle), then two blastodermic layers, with obliteration of the
cleavage -cavity and substitution of a primitive intestinal cavity

^ Not to Toe confounded with the original “germinal vesicle” of the parent-cell,
which long since disappeared.

334

GENERAL ORNITHOLOGY

PART II

(gastrula), is common to all animals which consist of more than
single cells, under various modifications and disguises ; the process
described is that occurring in meroblastic eggs which have a discoidal
cleavage and form a discogastrulad

What we have got now is a tread or germ consisting of a cir-
cular concavo-convex disc of two layers of blastoderm, resting by
its rim upon the great yellow ball of food-yelk, from which it is
separated by a cavity, as a watch-crystal from its face. All these
changes, up to completion of gastrulation, may go on before the egg is
laid, the tread of a perfectly fresh egg being already a multicellular

Fig. 112, — Further development of hen’s egg ; after Haeckel : A, the mulberry mass of
cleavage cells, h, same as seen on top in Fig. Ill, F, here viewed in profile in section, resting
upon n, the simply-shaded part of the figure, to represent conventionally the mass of food-
yelk. A, morula stage (as before) ; B, blastula stage, the mass of cells, &, forming the blasto-
derm, uplifted from the food-yelk, leaving the cleavage-cavity, s ; w, the thickened rim of the
germ-disc ; C, the blastnla in process of inversion, by which a layer of endoderin-cells, i, grow-
ing from periphery to centre, will apply itself to the layer of exoderm-cells, e, obliterating
the cleavage-cavity, s ; D, the discogastrula completed, by union of endoderm, i, with exo-
derm, e, leaving the primitive intestinal cavity, d, which is quite similar in appearance to
the cleavage cavity, s, but morphologically quite different.

discogastrula. Since the earlier stages of the embryo (cytula, morula,
blastula, and gastrula) are actually accomplished while the egg is
still in the body of the parent, the analogy of the oviduct to uterus,
etc., as well as its strict homology to the j^arts of a Mullerian duct
so named, is not so fanciful as some appear to think. The outer
of the two blastodermic layers is the ectoderm or epihlast, C or D, e ;
the inner is the endoderm or hjpoblast, L By multiplication of cells
between the two arises the mesoblast. The mesoblastic layer of
cells subsequently splits into two, of which the outer is the somato-
pleura, or body layer, the inner the splanchnopleura or visceral layer.
The two-layered germ has then become four-layered. Up to the

1 The so-called “ germ-vesicle ” of the holohlastic mammalian egg is subsequent
to gastrulation, not prior, and is therefore not a blastula proper.

SEC. IV

ANA TOMY OF BIRDS

335

time of formation of four layers, the cells are all alike, or only differ
slightly in size, colour, or consistency. Now, however, ensues that
marvellous process by which the indifferent cells of the blastodermic
layers are to become differentiated in form and specialised in function,
— a sort of division-of-labour system in the infant colony of cells,
by which some are to learn to move, others to digest, others to
procreate, others to think and feel, with corresponding modifica-
tions of form by which are generated the Osteamoehce, Myamcebce,
Neuramoebce, — the bone-cells, muscle-cells, nerve-cells, and all others
of the complex organism which is in a few days to come into being
from such simple beginnings. This of course opens up the whole
field of embryology, which we cannot here enter upon. I will only
add, that from the epiblast are derived the integument, and its in-
versions, as those of the eye and ear, and the brain and spinal
cord. From the hypoblast is derived the lining of the alimentary
canal and of its annexes and offsets, as liver, lungs, etc. The rest
of the embryo comes from the mesoblast, and most of it from the
somatopleural layer. The fissure between the two layers of the
mesoblast becomes the great pleuroperitoneal cavity.

In explaining the early embryo, I have closely followed the
great German morphologist Haeckel; and the illustrations are
from the same high source.

Incubation. — To induce the wonderful metamorphoses just
hinted at, it is only necessary to keep a bird’s egg at a pretty even
temperature of about 100° Fahr. Nearly all birds secure this result
by the process of incubation. In many cases the sun’s rays relieve
the parent of some part of the duty. In a few, the heat evolved
from vegetable ferment or decomposition is utilised for the same
purpose. This seems to be the case to some extent with grebes ;
but these incubate. “The exception to the rule of incubation is
given by the Megapodial birds of the Australasian Islands. A
huge mound of decaying vegetable matter is raised ; the eggs are
deposited vertically in a circle at a certain depth, near the summit,
and the chick is developed with the aid of the heat of fermentation.
The large size of the egg relates to affording a supply of material
suflScing for an unusually advanced state of development of the
chick at exclusion ; whereby it has strength to force its way to the
surface of the hatching-mound, with wings and feathers sufficiently
developed to enable it to take a short flight to the nearest branch
of an overshadowing tree ” (Owen). The period of incubation has
been ascertained with precision for few birds ; it is known to range
from ten days (perhaps less), as in case of the wren, to fifty or sixty
for the ostrich. The female is usually the sitter. Frequently both
sexes incubate in turn ; such unnatural care for the young by the
male is termed double monogamy. In most or all Ratitoi, in the

336

GENERAL ORNITHOLOGY

PART II

family Phalaropodidce, and some other Limicoline genera, the male
incubates. Most birds attend to their own eggs ; many cuckoos
(Cuculidm) and the species of Molothrus, are parasitical, laying in the
nests of other birds, which are thus forced to become foster-parents
of alien offspring, generally to the destruction of their own. This
seems to result from some peculiarity of the egg-laying process, which
does not permit several eggs to be incubated and hatched simultan-
eously. It is not so unusual among American cuckoos as generally
supposed. The degree of development to which birds attain in
the egg differs in AUrices and Prcecoces (see p. 131). They break
the shell by pecking at it, and struggling ; for the former operation
the bill is often tempered at the tip by a hard knob which is after-
ward absorbed. The necessity of providing a receptacle for eggs,
in which they may he incubated, results in nidification or nest-build-
ing ; and the extraordinary taste and ability many birds display in
this matter, as well as the wide range of their habitudes, furnishes
one of the most delightful departments of ornithology, called caliology
(Gr. KaXid^ kalia, a bird’s nest; see p. 81, note). Many birds
burrow in the ground ; others in trees ; the most beautiful and
elaborate nests are furnished by various members of the Oscines,
the weaver -birds of Africa (Ploceidce) probably taking the lead.
The male sometimes constructs his own “ nest ” apart from that in
which the female incubates. “Certain conirostral Cantor es still
practise in the undisturbed wilds of Australia the formation of
marriage-bowers distinct from the later-formed nesting-place. The
satin bower-bird {Ptilonorhynclius holosericeus)^ and the pink-necked
bower-bird {Chlamydodera maculata), are remarkable for their con-
struction on the ground of avenues, overarched by long twigs or
grass-stems, the entry and exit of which are adorned by pearly
shells, bright-coloured feathers, bleached bones, and other decorative
materials, which are brought in profusion by the male, and vari-
ously arranged to attract, as it would seem, the female by the show
of a handsome establishment ” (Owen). The extraordinary nests of
the Crotophaga are used in common by a colony of the birds. “ Edible
birds’ nests,” constructed by swifts of the genus Collocalia^ consist
chiefly of seaweeds and inspissated saliva. Perhaps the most re-
markable of all the receptacles of eggs is that which the penguin
makes of its own body, the egg being taken in a sort of pouch
formed by the integument of the belly, something like that of a
marsupial mammal.

INDEX

Abdomen, 93, 137, 140
Acetabulum, 176, 219
Adrenals, 69, 322
iEgitlioguathism, 254
^pyornis Maximus, 99
Age, recognition of, 70
Air-gun, 7

Ala spuria (Alula), 162
Alca impennis, 99

Alimentary annexes, 317 ; canal, 309

Allantois, 92, 292

Altricial, 131

Ambiens, 287

Ammunition, 4, 5, 7

Amnion, 92

Amoebiform animals, 285

Amphicoelous, 203

Analogy, 103

Anomalogonatous, 289

Antebrachium, (a) ulna, (b) radius, 158

Anthrenus, 84

Antitrochanter, 219

Aortic arches, 93

Apophyses, 199, 202, 303

Apteria (spaces), 130

Aptosochromatism, 132

Aqueous humour, 265, 272

Arachnoid, 260

Archaeopteryx, 96, and illustration

Archetypical characters, 116, and note

Arsenical soap, 39 note

Arterial system, 290

Articulation, 199

Arytenoid, 301

Astragalus, 94

Atmosteon, 249

Attypical characters, 116 note

Audition, 273

Auk, great, 99

Auriculars, 136, 143, 274

Aves ,92, 93 ; position in animate nature,

94 ; class or an order or sxib-class, 94 ;
geological succession, 96 ; archaeo-
pteryx oldest known, 96

Z

Barba, 126
Barbicels [cilia), 126
Barbules, 126
Basitemporal bone, 229
Beards, 145

Bicarotidinae normales, aves, 294 ; abnor-
males, 294

Bill [rostrum), 136, 146 ; (1) land birds,
fissirostral, (2) tenuirostral, (3) denti-
rostral, (4) conirostral, 148 ; (5) longi-
rostral, (6) pressirostral,(7) cultrirostral,
(8)lamellirostral, 149 ; covering of, 150 ;
cere, 150 ; caducous or deciduous, 151 ;
rhamphotheca, [a) rhinotheca, [b) gna-
thotheca, 151

Binomial nomenclature, 121
Biogenation, 284
Bird-lime, 7

Birds, how to trap, 7 ; how to net, 7 ; how
and where to seek, 15 ; quantity re-
quired, 19 ; how to approach, 23 ; how
to find shot, 25 ; how to kill wounded,
25 ; how to handle bleeding, 26 ; how
to carry, 27 ; how to mount, 60 ;
definition of, 92 ; classification of, 99 ;
taxonomy, 100 ; reason for morpho-
logical classification, 104
Birdskins, how to prepare, 38-42 ; (1)
instruments, (2) materials, 38 ; [a) for
stuffing, [b) for preserving, 39 ; (c) for
cleansing, 40 ; [d) for wrapping, 41 ;
how to make, 42 ; [a) regular process,
42-51 ; position of bird, 43 ; [b) special
processes, 52-68 ; with reference to size,
52 ; shape, 54 ; thin skin, loose plumage,
55 ; fatness, 56 ; bloodstains, etc., 60 ;
restoration, 71
Blastoderm, 333
Blastula (germ vesicle), 333
Blood, 289 ; serum, coagulum, plasma,
290

Bloodstains, 56

Blowing eggs, 7 6 ; method of strengthen-
ing egg before, 79 note

338

INDEX

Blow-gun, 6

Body or Trunk (corpus), 137
Bones — basioccipital, 93 ; quadrate, 93 ;
tarsal, 93 ; carpal, 94 ; interclavicular,
94 ; wish-bone or merrythought, 94 ;
sternum, 94 ; ilia, 94 ; femur, 94 ;
astragalus, 94 ; tibia, 94 ; fibula, 94 ;
metatarsus, 94 ; pneumaticity of, 199
Brain {cerebrum) — (a) prosencephalon,
(&) mesencephalon, (c) opisthencepha-
lon, {d) diencephalon, (e) epencepha-
lon, 258 ; general characters of, 260
Branchial bones — (1) cerato-, (2) epi-,
(3) basi-, 226

Breast-bone {sternum), 73 ; illustration,
137

Breech-loader v. Muzzle-loader, 4
Breech-loading j^istol, 6
Bursa Fabricii, 317

Cabinets, 84

Cseca — (a) coli, {b) umbilical or vitelline,
316

Calamus, 125
Caliology, 336

Camptolemus labradorius, 99
Cane-gun, 6

Canthus — {a) anterior, {b) posterior, 143
Capsule (calyx), 327
Cardiac, 315
Carpals, 94

Carpus — {a) scapholunare, (5) cuneiforme,
158

Cartridges, 5
Caruncles, 145
Cases, 84

Caudal (Coccygeal) vertebrae, 209
Centra, 203
Cerebellum, 259
Cerebrum (brain), 258
Cervical, (1) atlas, (2) axis, 206 ; verte-
brae, 136, 204

Chalaziferous membrane, 329
Chest (thorax), 137
Choroid membrane, 270 ; plexus, 260
Chyle, 295

Cicatricle (tread), 327
Cilia (barbicels), 126
Class, 110

Classification — morjihological or homo-
logical, 101, 102 ; physiological or
analogical, 102 ; reason for morpho-
logical, 102 ; dangers of morphologi-
cal, 105 ; zoological, 105
Clavicles { furculum), 217
Claviculae (collar bones), 137
Claws, 168, 195
Cleavage cell in egg, 332
Clinoid wall, 226

Clitoris, 329
Cloaca, 93, 317
Clutch of eggs, 80
Coagulum, 290
Coccyx, 169

Coccygeal (caudal) vertebrae, 209
Cochlea, 224
Cochlear cavity, 275
Cold-taking, 29
Collar-bones {elaviculse), 137
Columella auris (stapes), 275
Combs, 145
Commissure, 155
Conarium (or pineal body), 259
Condyles, 93, 199
Conspecies {subspecies), 121
Contour feathers, 128
Contour of bird, 134
Coracoid, 73, 137, 216
Coracostea, 214
Cornea, 265

Corniculum laryngis, 301

Corona (vertex), 142

Corpus Callosum, 93

Costal process, 211, 212

Cotyloid (coxal) cavity, 219

Cranial bones, 237 ; nerves, 261

Crests, 145

Cricoid, 301

Crissum, 140, 170

Crotaphite depression, 232

Crus — {a) tibia ; (5) fibula, 177, 186

Crystalline lens, 265, 271

Culmen, mandibular, 152

Cupola, 279

Cytula, 332

Darwin on plumage, 133
Decomposition, 59
Dentine, 198
Dermestes, 82
Dermestidse, 82
Desmognathism, 253
Diapophysis, 202
Diaphragm, 93
Didusineptus, 99
Diencephalon, 258
Dinornithes, 99
Discogastrula, 333
Dodo, 99
Dogs, 14

Dorsal part of body, 138
Dorsal (thoracic) vertebrae, 206
Dorso-lumbar, 208
Double-barrelled shot gun, 3
Double monogamy, 335
Dromseognathism, 249
Duodenum, 316
Dura Mater, 260

INDEX

339

Dyuamamoeba, 318

Ecdtsis (moult), 131
Ectoderm (epiblast), 334
Eggs, instruments for preparing (blowing),
76 ; method of numbering, 80; clutch
of, 80

Elseodochon (uropygial gland), 129
Embryological characters, 107
Embryology, 320

Endoderm (hypoblast), 334 ; cells in an
egg, 333
Endolymph, 282
Endysis, 131
Enteramoeba, 310
Epencephalon, 258
Epiblast (ectoderm), 334
Epidermic, 123
Epiclidium, 217
Epigastrium, 140
Epiphyses, 93, 199
Equipments — miscellaneous, 8
Equivalence of groups. 111
Ethmoid, 236 ; plate — aliethmoid, ali-
septal, alinasal, 226
Etypical characters, 116 note
Eustachian tube, 234
Exoskeletal, 123
Extensors, 161
Eye, 264
Eye-baby, 271

Facial bones (or Vomer), 237

Fallopian ISTerviduct, 278

Family, 110

Fascife, 285

Fat, 56

Fatigue, 31

Feathers, outgrowths or appendages, not
tegumentary expansion, 123 ; (A)

development, 124 ; structure, main
stem or scape {scapus), supplement-
ary stem {hyporhacMs), webs or
vanes [vexillum), scape, {a) barrel or
quill {calamus), 125 ; (5) shaft proper
{rhachis), barb {harha), barbules, barbi-
cels [cilia], booklets [hamuli], 126 ;
(B) kinds of — (1) contour [pennce or
plumce], (2) down [plumulce], (3) semi-
plumes [semiplumce], (4) filoplumes
[filoplumoe], 128 ; (5) powder down
[pulviplumes], 129 ; (C) of the wing —
flight [remiges], 162, 164 ; coverts
[tectrices], alula, or ala spuria, 162
Feathery structure, types of, (1) plu-
mous or pennaceous (feathery), (2)
plumulaceous (downy), 127 ; (3) filo-
plumaceous (hairy), 128
Female organs of generation, 324
Femur, 94

Fenestra, 213 ; ovalis, 227, 275 ;

rotunda, 275
Fibula, 94

Field-work, miscellaneous, 15
Filoplumae, 128
Fixtures, 41
Flaps, 145
Flexors, 161
Flocculus, 277
Fontanelle, 213
Food-yelk, 327

Foot, (A) bones of, thigh or femur, 175 ;
acetabulum, 176 ; crus, [a] tibia, (6)
fibula, heel or suffrago, 177 ; digits or
toes, phalanges, internodes, mechan-
ism of, 180 ; plumage of, 182 ; length
of, horny integument [podotheca],1^2> ;
[a] scales [scutella], 184 ; (6) plates or
reticulations, (c) granulation or rugose,
[d] scabrous or scarious, [e] cancellated,
(/) serrated, [g] holothecal, [h] schizo-
thecal,185; (B) types of — (l)insessorial
or perching, 190 ; (2) cursorial or
grallatorial (walking or wading), 193 ;
(3) natatorial or swimming, 194 ; [a]
palmate, (6) lobate, semipalmation,
193

Foramen, 205 ; ovale, 224, 236 ; mag-
num, 230 ; optic, 236 ; olfactory, 237;
pneumatic, 246

Forms, [a] generalised, [h] specialised,
117

Frontals, 231

Furculum (merrythought), 136 ; (clavi-
cles), 217
Fusiform, 134

Ganglia, 257
Gastornis parisiensis, 98
Gastreeum, [a] breast [pectus), [h] belly
(abdomen), 140
Gastrula, 333
Geant, 99

Gena (malar region), 144
Generation, female organs of, 324
Geniohyoid muscle, 312
Genital gland, 319
Genus, 110
Germination, 331
Germ-vesicle (blastula), 333
Germ-yelk, 327
Gigerium, 314
Gizzard [gigerium], 314
Glenoid cavity, 217
Glossopharyngeal nerve, 284
Glottidian fissure, 301
Gonys, 246

Graafian follicle (ovisac), 326 ; gula, 144

Gullet (cEsophagus), 136, 313

Gun, double-barrelled shot, 3 ; breech-

340

INDEX

loader v. muzzle - loader, 4 ; single
barrel, 6 ; cane, 6 ; blow, 6 ; air, 7 ;
how to carry, 9 ; how to clean, 10 ;
how to load, 11 ; how to shoot, 12 ;
accidents from, 29
Gustation, 283
Guttur, 141

H^emapophysis, 203
Haematamoeba cruentata, 291
Haematothermal, 291
Halluae, characteristics of, 190
Hamuli (hooklets), 126
Harderian gland, 266
Harpagornis, 99

Haunch bone (os innominatum), 138, 219
Head {Ca'put\ 135
Heart, 291

Hesperornis, 96, illustration on
Heterocoelous, 204
Homalogonatous, 289
Homology, 103
Hooklets {hamuli), 126
Humero-scapulare, 215
Humerus, 157
Hunger, 31

Hyoid bone, ceratohyal, basihyal, 225,
247 ; branchials, 247, 248
Hypapophysis, 203
Hypoblast (endoderm), 334
Hypoclidian, 217
Hyporhachis, 125

ICHTHYOPSIDA, 92

Ichthyornis, 96, illustration 98

-idee as a termination, 120

Ileum, 316

Ilium, 94, 138, 219

-inoe as a termination, 120

Incubation, 335

Incus, 276

Infundibulum, 279, 328
Ingersoll (Mr. Ernest), on nest-collect-
ing, 81 note

Insect pests, 82 ; how to destroy, 85
Intermaxillary bones, 242
Interorbital septum, 226
Intestine, 315

Intestinal cavity in an egg, 333
Iris, 265, 271
Ischium, 138, 220
Isthmus of the oviduct, 328

Jejunum, 316

Jugal (quadratojugal) bone, 240
Jugulum, 141

Kidneys, 69, 93, 321

Labels, 34

Labrador Duck, 99

Lacrymal bones, 244 ; palatinum, os, 245
Lacteals, 295

Ijsevo-carotidinae aves, 294
Lamina spiralis, 279
Larynx, 300
Leguata gigantea, 99
Leucocytes, 291
Lingula, 224
Liver, 318
Lobes, 145
Lophosteon, 212
Lore, 143

Lumbar vertebrae, 207
Lungs, 296
Lymph, 289
Lymphatics, 295

MalaPv region {gena), 144
Malleus, 276
Mammalia, 92

Mandible, {a) lower, tomium, 152 ; (6)
rami, (c) gonys, 152 ; {d) upper,
culmen ; (e) maxillary tomium, 153 ;
nostrils, 153 ; gape, 155
Mandibles, 147, 224, 245 ; (1) epigna-
thous, (2) hypognathous, (3) para-
gnathous, 148 ; (4) metagnathous,
148

Mantle {straguluvi), 140
Manubrium (or rostrum), 214
Marriage bowers, 336
Marsupium pecten, 272
Maxillopalatine, 240 ; (sub-ocular) bone,
224

Maxillary bone, 240
Measuring of birds, 37 ; length ; extent;
length of wing, of tail, of bill, tarsus,
37 ; toes, claws, head, 38
Meatus, 234 ; auditorius, 143 ; {a) ex-
ternus, 274 ; (&) internus, 278
Meckelian cartilage, 225
Medulla oblongata, 260
Membrana tympani, 228
Meninges, (1) pia mater, (2) arachnoid,
(3) dura mater, 260
Mentum, 144
Meroblastic, 327

Merrythought {furculum), 73, 136

Mesencephalon, 258

Mesenteries, 310

Mesoblast, 334

Mesometry, 328

Metacarpus, 158

Metatarsus, 94

Metencephalon, 258

Metosteon, 213

Metovum, 327

Miasm, 29

Moas, 99

INDEX

341

Monerula, 332
Monogamy, double, 335
Morphology, 101
Morsus diaboli, 328
Morula, 333

Moult {ecdysis), number of, 131
Mouth, 310
Mullerian duct, 319
Mummification, 72
Muscular tissue, 285
Mutilation, 58

Muzzle-loader v. breech-loader, 4
Myamoebse, 285, 335
Myelencephalon (or Metencephalon), 260
Mylohyoid muscle, 311

Nasal fossa (groove), 153 ; bones, 243 ;

holorhinal, schizorhinal, 244
Neck {collum), 136 ; {a) nucha, (5)

gula, (c) cervical, 141
Nests, advantages of collecting, 81 note
Nestor productus, 99
Netting birds, 7

Neuramceba, 335 ; {a) cinerea, (5)

Candida, 257
Neurapophysis, 202
Nictitating membrane, 266, 267
Nidification (nest-building), 336
Nomenclature, {a) binomial, 119, 120 ;

(6) trinomial or trionymic, 121 note
Non-vascular capillary, 290
Notaeum, (a) back {dorsum), 138, 139 ;
{h) rump {uropygium), (c) scapulare,
(d) interscapulare, 139
Note-book, best kind of, 34
Notochord, 224

Observations, record of, 33

Occipital bone, 230

Occiput, 142

Odontolcee, 97

Odontotormse, 97

CEsophagus (gullet), 136, 313

Oil, gland for, 129

Olecranon, 158

Olfaction, 263

Ontogeny, 108

Oology, 75, 320

Oophron masculinum, 323

Opisthencephalon, 258

Opisthocoelous, 204

Optic nerve, 272

Orbit of head, 143

Orbitonasal septum, 236

Order, 110

Ornithichnites, 96

Ornitholite, 96

Ornithology, definition, 91

Os innominatum (haunch-bone), 219

Ossiculum auditus, 276

Osteamcebae, 221 note, 335

Osteine, 198

Osteology, 198

Osteoses, 199

Otocrane, 277

Otolith, 282

Ovaries, 69, 93

Oviduct, 69, 319

Oviparous, 92

Oviposition, 330

Ovisac (Graafian follicle), 326

Ovoviviparous, 92

Ovulation, 326

Ovum, 326

Palatal, or palatine, bones, 241 ; struc-
ture, types of, 249
Palpebral layer, 268
Pancreas, 317
Parachordal cartilage, 224
Parasphenoid bone, 229

*pQ Yi-j p+o 1 Qj 0^1

Parker (Mr. W. K.), on the skull, 223
Pectorales, 286

Pectoral arch, (1) scapula, (2) coracoid
(together are scapular arch), (3) cla-
vicles, 215
Pectus, 140
Pedicel, 327

Pelvis, 137 ; (a) ilium, (6) ischium, (c)
pubis, 218
Perilymph, 282
Periotic bones, 232
Pessulus, 303
Petit, canal of, 272
Pezophaps solitarius, 99
Phalanges, number of, 159, 180, 188
Phylogeny, 108
Phylum, 101

Physiological adaptations, (a) pinnati-
pedes, 102
Pia mater, 260
Pileum (crown), 142, 143
Pineal body (or conarium), 259
Pinnatipedes, 102
Pistol, breech-loading, 6
Pituitary space, 224
Plasma, 290
Pleurapophysis, 203
Pleurosteon, 212

Plumage {ptilosis), 123-134; changes,
133

Plumulse, 128
Pneumaticity, 199
Pneumatocysts, 296
Pocket lens, necessity of, 41
Postfrontal process, 231
Postzygapophysis, 202
Powder, 7
Prcecocial, 131

342

INDEX

Premaxillary bones, 242
Prenasal cartilage, 227
Prezygapophysis, 202
Procoelous, 204
Procoracoid, 215
Prosencephalon, 258
Prototype, 115
Protovnm, 327
Protozoa, 108
Proventriculus, 314
Psilopsedic, 131

Pterygoid bones, 240 ; process, 242
Pterygopalatine rod, 229
Pterylse (tracts), 129
Pterylography, 130
Ptilopaedic, 131
Ptilosis (plumage), 123
Ptinus, 82

Pubis, 138 ; (a) epipubis, (&) propubis,
220

Pulviplumse, 129
Pupil, 271

Putaminis membrana, 330
Pygastyle, 169, 210
Pylorus, 315

Quadrate bone, 93, 239
Quadratojugal (jugal) bone, 240

Eectrices (rudders), 170

Eemiges, 162, 164 ; (a) primaries, 166 ;

(d) secondaries, 167
Eeptilia, 92

Eespiration, organs of, 295
Eetina, 272
Ehachis, 126
Ehinencephalon,, 295
Eibs, (a) fixed, (&) fioating, (c) sacral,
210

Eictal bristles {vibrissee\ 146
Eictus, 155
Eigor mortis, 59
Eima glottidis, 301
Eostrum (manubrium), 214
Eudders {rectrices), 176

Sacral plexus, 208
Sacrocostal, 211
Sacrum, 138, 207
Saurognathism, 255
Sauropsida, 92

Scapula, 137, 217 ; accessoria, 162, 215

Scapular arch, 73, 137

Scapus, 125

Schizognathism, 252

Scrotum, 69

Semen, 323

Semilunar membrane, 305
Semipalination, 193
Semiplumse, 128

Sensorimotor, 258
Serum, 290

Sex, recognition of, 68 ; {a) testicles,
(6) ovaries, (c) scrotum, {d) sperm-duct,
(e) oviduct, (/) kidneys, {g) adrenals,
69

Shoot, how to, 12 ; times to, 18
Shot, 4, 5, 7

Single-barrel gun, 6 ; single-emarginate,
single-notched, 213
Sinus rhomboidalis, 261
Sitodrepa, 82

Skeleton (A) — (a) endoskeleton, (&) exo-
skeleton, (o) scleroskeleton, 198 ;
(B) — Axial, {a) neural arch, 200 ; (b)
haemal arch, 201

Skull, 222, illustrations, 222, 223

Smell, sense of (olfaction), 263

Solitaire, 99

Somatopleura, 334

Spaces {apteria), 130

Species, 110

Spermatozoa, 323

Sperm-duct, 69

Sphenoid, 234

Sphenotic bone, 231

Sphincter vaginae, 329

Spinal cord, 261 ; nerves, 262

Splanchnopleura, 334

Spur {calcar), 168, 196

Spurious, two uses of, (c) tertiaries, 167

Squamosal, 232

Stapedial, 227, 228

Stapes {columella auris), 227, 275, 276
Stenson, duct of, 311
Sternal {a) ratite, (&) carinate, 212
Stigma of an egg, 327
Stimulation, 31
Stomach, examination of, 70
Sternum (or breast -bone), preservation
of, 73 and illustration ; avian, 211
Subgenus, 122

Subocular (or maxillopalatine) bar, 224
Subspecies {conspecies), 121
Suffrage, 177
Suprarenal capsules, 322
Suspensorium, 225
Suture, 199

Sympathetic system, 263
Synovia, 199
Syrinx, 301

Taction, 283

Tail, (1) bony basis, (2) . coccyx, (3)
vomer, (4) pygostyle, 169 ; tail-coverts
{tectrices), 169 ; shape of, 173
Tarsal bone, 93
Tarso-metatarsus, 94
Tarsus, 186

Taxonomy, 100 ; (1) morphological, 101

INDEX

343

Tectrices, 162
Teleotype, 115
Tensor patagii, 287
Testicles, 69

Testis or testicle, 69, 93, 322
Thalamencephalon, 259
Thigli {femur), 175
Thoracic (dorsal) vertebrae, 206
Thorax, 93, 210
Thyroid, 301
Tibia, 94

Times to shoot, (a) time of year, (5) time
of day, 18 ; (c) weather, 19
Tincae, os, 329

Tinea, {a) flavifrontella, (5) tapetzella,
82

Tineidae, 82

Toes {digiii), number of, 187 ; didactyl,
tridactyl, tetradactyl, zygodactyl, posi-
tion of, 189

Tomium, 155 ; mandibular, 152 ; max-
illary, 153
Tongue, 312
Tonicity, 285
Topography of a bird, 134
Trabeculae, 224

Trachea (windpipe), 74, 75 illustration,
298

Tracts (pterylae), 129 ; (1) spinal or dorsal
{joteryla spinalis), (2) humeral, (3)
femoral, (4) ventral, 130 ; (5) head,
(6) wing, (7) tail, (8) leg, 131
Trapping birds, 7
Tread (cicatricle), 327
Trinomial nomenclature, 121
Tympanum, 274

Types of structure, (<x) typical, (5) sub-
typical, (c) aberrant, 114 (c^) tele-

otypic, (e) prototypic, 115

Uncinatum, os, 245
Ureters, 317
Urinary organs, 318
Urogenital sinus, 317
Uropygial gland (elaeodochon), 129
Urosacrals, 218

Urosteon, 214

Valuation of characters, 112
Vasa deferentia, 319
Vascular capillary, 290
Venous system, 290
Venter, 140

Ventral part of body, 138
Ventri cuius bulbosus, 314
Vertebra, 203
Vertebrarterial canal, 205
Vertebrata, {a) aves, (5) reptilia, 92
Vesiculae seminales, 322
Vexillum, 125
Vibrissae, 146
Vision, 264

Vitellus (yelk), 326 ; segmentation of,
332

Vitreous humour, 272
Viviparous, 92
Vomer, 169, 237

Wads, 8

Wattles, 145

Wet preparations, 72

Windpipe (trachea), 74, 75 illustroMon

Wing-bones, mechanism of, 162

Wing-coverts, 162

Wings, 155 ; humerus, 157 ; cubit or
antebrachium, {a) ulna, {h) radius,
olecranon, carpus, {a) scapholunare,
(&) cuneiforme, metacarpusj 158 ;
length of, 168
Wolffian bodies, 93, 318

Xiphoid process, 213

Yelk {vitellus), 326 ; {a) food, {h) germ,
327

Zona pellucida, 326 ,

Zoological characters, 107 ; groups,
class, order, family, genus, species,
110

Zygoma, 230
Zygapophysis, 202

THE END

'dinbur^h