..v*'.-

• ■ f^::

FOR THE PEOPLE

FOR EDVCATION

FOR SCIENCE

LIBRARY

OF

THE AMERICAN MUSEUM

OF

NATURAL HISTORY

'O.

THE

MYOLOGY OF THE EAVEN ^'

{Corviis cor ax sinuatus).

A GUIDE TO THE STUDY OF THE MUSCULAR SYSTEM
IN BIRDS.

E. W. SHUFELDT.

Eontron:
MACMILLAN AND CO.

AND NEW YORK.
1890.

The Right of Translation and Reproduction is Reserved.

,^.b^jn(^i'-'^

RicnABD Clay and Sons, Limited,

LONDON AND BUNGAY.

PREFACE.

There has not yet been published, so far as the writer
is aware, any work devoted to a complete account of the
muscles of any single species of the Class Aves, — a work
fully and practically illustrated, and one that would
prove itself to be of service to those either engaged in
the general study of the morphology of vertebrates, or
to those special students who may be investigating the
myology of birds. At the present time, — when the
study of the structure of animals is becoming far more
general, as one of the most efficient aids to observation
and mental training, than it was so considered a number
of years ago, — books of the class which your author
has here endeavoured to produce come to be very
useful. Birds stand among the most easily procurable
subjects for the use of the demonstrator at the laboratory,
or for the student to employ in his own researches at
home as illustrative of certain parts of his course in
biology. And it was to fill this so important a gap, as
the lack of a suitable volume devoted to the muscular
system of birds, that the writer undertook an exhaustive
study of the muscles of the Raven [Corviis corax

PREFACE.

sinuatus) ; and the work here placed before the reader,
with its eighty and odd figures, is the outcome of those

investigations.

With regard to my nomenclature of the muscles T
have here descril)ed and figured, I have been guided by
such brief chapters as are devoted to this sul)ject and
which appear in the published scientific memoirs of
such excellent authorities as Owen, Carus, A. Milne-
Edwards, Huxley, Garrod, Forbes, Selenka, Coucs, Fiir-
bringer, and Gadow ; but when I have been in doubt,
and these authorities failed to assist me, I have en-
deavoured to bestow upon the muscle a suitable name.
The student must bear in mind, too, that many other
birds possess muscles which are not to be found in the
Raven ; in some special cases I have alluded to these.
On the other hand, the muscles, even in the represen-
tatives of the same species, may vary in certain in-
dividuals to some extent. This fact has lono; been
appreciated by anthropotomists.

Gadow's work upon the muscles of birds, which
appeared in Bronn's Thier-Iieichs, is an admirable con-
tribution to the general subject ; but it is by no means
a work that meets the general want, and possesses the
disadvantages of being but meagrely illustrated, and of
having appeared in German, in a work of limited cir-
culation. Notwithstanding this, I am quite sure my
reader will feel grateful for my having incorporated
Gadow's synonymy in footnotes in the present volume,
as they cannot fail to be anythiug but the most useful
adjunct to a guide to avian myology.

In arranging and grouping the elements of this

PREFACE. vii

system, it is to be hoped that the writer's plan will be
found to be a convenient and helpful one : it was his
chief aim throughout, next to accuracy and clearness of
description, to make it so.

In choosing the Raven for our subject, it was done in
view of the fact that it is a large representative of a
very numerous and cosmopolitan family of birds, the
CorvidcB ; so that, in almost any part of the world, a
variety of birds become available whose muscular sys-
tems can be studied by the aid of the present volume.
It is hardly necessary to add that C^rows of all descrip-
tions, Jays, Orioles, and a host of others, all fall within
this category. It has its advantage, too, for the teacher
and the student at the biological laboratory ; for the
former can use as his subject the larger and more advan-
tageous specimens, as the Ravens or Crows, while the
latter can confirm the instructions of the former, at
home, upon any of the smaller varieties of the Corvidce,
such as the Jays or Rooks.

As his investigations in the myology of vertebrates
jDrogress, three lines for improvement, in so far as our
knowledge of the muscular system of birds is concerned,
will force themselves upon the student. In the first
place, we still remain very ignorant of the details of this
system in a great many important types of birds ;
secondly, an ever-pressing demand is evident, to fix the
homologies of muscles in the Vertebrata, and conse-
quently to bring so far-reaching a knowledge of this
department of research to our assistance as to be enabled
to give the same name to the same muscles, accurately,
throughout the vertebrate series ; finally, a simple.

viii PREFACE.

scientific, and euphonious nomenclature is very much
to be desired. As an index of our present status with
respect to our knowledge of the muscles of birds, it is
hoped that the volume here offered Avill faithfully repre-
sent it, Init its writer trusts that in future works he
may lend his assistance to the improvement of all the
lines above indicated. To this end, he will be thankful
for any suggestions that may be useful should the
present volume go to another edition, and good criticism
of the same will be welcomed from any quarter.
Already, the question of neuro-myology, or the nomen-
clature of muscles based upon tlieir nerve-supply, has
made considerable progress in modifying our views in
that direction, and although I am convinced that it
by no means always furnishes an invariable guide to
follow, still its great usefulness is not to be ignored.
With no little interest I have read the " Neuro-myology "
of Coues and Shute [New York Medical Record, July,
1887), and am compelled to admire the masterly manner
in which they have instilled new life into an old theme.
Unfortunately, the same muscle as represented in
diff"erent types of the vertebrate series, is not always
served by the sa7ne nerve, and this w^ould stand in the
w^ay of a general grouping and a universal neuro-
myological nomenclature upon any such a basis. To
those of my readers who are familiar with German, the
best works 1 can recommend to be consulted in the
present connection are the very excellent treatises of
Selenka and Gadow in Bronn's Klassen des Thier-Reichs;
and that superb monument to avian morphology, the
Untersuchumjen zur Morphologic und Systematik der

PREFACE. ix

Vogel of Max Fiirbringer. To Professor Gadow I am
indebted for a great many titles of works which I have
incorporated in the bibliography which is found at the
end of this volume ; still, I have personally examined
the majority of books there enumerated.

English and American literature, as I have already
stated, furnishes us with no good handbook to the
subject.

Frequently, an author, as he closes his prefatory
remarks, finds that he has a host of helpful friends to
whom his thanks are due for assistance rendered : it
proves to be the exception in the present instance, for
all of the material used was collected by myself ; all the
dissections were made by myself ; all the descriptions
are in my own handwriting ; and the drawings were all
drawn directly from the dissections by myself. Still it
gives me pleasure to remember here my friend Dr. J. L.
AVortman, the biologist of the United States Army
Medical Museum, who, several years after this volume
was written, carefully passed over wnth me the mus-
culature of the shoulder-joint in birds, and verified my
dissections upon the Eaven, comparing them with
similar studies he was at the time engaged upon in the
Raptores. Further it gives me pleasure to express my
thanks to Dr. G. Brown Goode, of the United States
National Museum, for having upon a number of
occasions furnished me with the means of preserving
my somewhat extensive material, and that, too, when
my labours upon the present volume were being
prosecuted in New Mexico, between two and three
thousand miles from civilization, the libraries, and the

X PREFACE.

museums ; and tliat at a time when the operations of
a short-siglited and destructive policy which influences
our Government in its attitude toward scientific inves-
tigators came quite near placing the researches herein
tendered to my readers and collaborators in anatomy,
iDcyond the pale of publication.

• R. W. S.

Takoma, D.C.

Anijuat 20, 1889.

CONTENTS.

PAGE

PREFACE V

LIST OF ILLUSTRATIONS Xui

INTRODUCTION 1

THE DERMAL MUSCLES 3

THE MUSCLES OF THE HEAD 15

THE MUSCLES OF THE TONGUE 23

THE MUSCLES OF THE AIR PASSAGES 43

THE MUSCLES OF THE EYE AND EAR 53

THE MUSCULATURE OF THE EAR 62

THE MUSCLES OF THE UPPER EXTREMITY 66

THE MUSCLES OF THE LOWER EXTREMITY 155

THE MUSCULATURE OF THE TRUNK 258

BIBLIOGRAPHY^ 319

INDEX , 329

LIST OF ILLUSTEATIONS.

[Note : — All the figures illustrating this woi-k were drawn by the
author, and with but few exceptions, where they have been
copied from the drawings of other anatomists, were made
directly from the recent dissections, and in the vast majoi'ity
of cases the size of life.]

FIG. rAGK

1. Side view of the skull of the Raven, showing points of

origin and insertion of muscles 7

2. Superior aspect of skull of the Raven, showing points of

origin and insertion of muscles 11

3. Showing points of attachment of muscles on the basal

aspect of the skull in the Raven 17

4. The posterior aspect of the skull in a Raven, showing

origin and insertion of the muscles 19

5. Side view of the mandible in Corvus corax sinuatus, show-

ing the areas of muscular origin and insertion .... 21

6. Corvus corax sinuatus, the mandible from beneath, showing

points of attachment of muscles 23

7. Corvus corax sinuatus, dissection of muscles of the head,

seen upon right lateral view 28

xiv LIST OF ILLUSTRATIONS.

no. rAGF.

8, Corviis corax sinuatus, oblique view of bones of shoulder-

girdle, showing areas of attachment for muscles ... 31

9. Corvus corax sinuatus, the hyoid arches from beneath,

showing points of attachment of muscles 31

10. Corvus corax sinuatus, superior view of bones of the upper

larynx, showing points of attachment of muscles ... 31

11. Corvus corax sinuatus, tbe muscles of the head, seen from

beneath 32

12. The tongue and salivary glands of a Woodpecker (after

Owen) • 44

13. Front view of the lower larynx of a Raven, showing

muscles (after Owen) 44

14. Side view of the lower larynx of a Raven, showing mviscles

(after Owen) 44

15. Lower larynx of a Parrot (after Owen) 44

IG. Muscles of organ of hearing in an Owl (after Owen) . . 44

17. Corvus corax sinuatus, its hyoid arches from beneath,

showing points of muscular attachment 46

1 8. Corvus corax sinuatus, its hyoid arches from above, show^-

ing points of miiscular attachment 16

19. Corvus corax sinuatus, the musculature of its windpipe . . 50

20. Corvus corax sinuatus, the head from above, with roof of

orbit removed 54

21. Mesial aspect of an eye of a Goose (after Owen) .... 55

22. Corvus corax sinuatus, side view of skull to show origin of

eye-muscles in the orbit 56

LIST OF ILLUSTRATIONS. xv

FIG. PAGE

23. Corviis co7'ax siviuatus, muscles at the back of the eye, dis-

sected , 60

24. Corvus cor ax sinuatus, right lateral view of the skeleton

of the trunk, showino- areas of muscular attachment . 68

25. Corvus corax sinuatus, the sternum from in front, showing

areas of mviscular attachment 71

26. Corvus corax sinuatus, superficial muscles, antei-ior aspect,

of the upper extremity, as far as the elbow 77

27. Corvus corax sinuatus, anconal aspect of right humerus,

showing where the muscles ai'ise and are inserted ... 83

28. Corvus corax sinuatus, proximal view of head of humerus,

to show muscular attachments 83

29. Corvus corax sinuatus, palmar aspect of humerus, to show

points of muscular attachment 83

30. Corvus corax sinuatus, radial aspect of humerus, to show

points of muscular attachment 83

31. Corvus corax sinuatus, ulnar aspect of humerus, to show

areas of attachment of muscles . . . . , 83

32. Corvus corax sinuatus, anterior aspect of some of the

muscles of the upper extremity 89

33. Corvus corax sinuatus, superficial muscles of the upper ex-

tremity in the region of shoulder and dorsum .... 97

34. Corvus corax sinuatus, oblique left lateral view of the

second layer of muscles of the upper extremity, being
those of the dorsum and arm 103

35. Corvus corax sinuatus, left oblique view of the body, dis-

sected to show the deep muscles about the shoulder . . 107

xvi LIST OF ILLUSTRATIONS.

FIG. PAGE

35 his. Axillary muscles .of the side of Gallinula chloropus

(after Garrod) 110

35 ter. Outer aspect of the axillary muscles of the right arm
in. a specimen of the Mourning Dove {Zenaidura
tnacroura) 110

35 quat. The plucked body of a bird, drawn to show where

to make the incision to expose the patagial muscles . . 112

35 quin. Muscles of the patagium in Ictertis vulgaris (after

Garrod) 113

35 sex. Muscles of the patagium in Progne subis 114

36. Corvus corax sinuatus, bones of forearm and hand, showing

muscular attachments 121

37. Corvus corax sinuatus, radial aspect of left ulna and radius,

showing where muscles are attached 126

38. Corvus corax sinuatus, outer aspect of left forearm and

pinion, showing the position, origin, and insertion of

the superficial layer of muscles 131

39. Corvus corax sinuatus, inner aspect of left forearm and

pinion, showing the position, origin, and insertion of

the superficial layer of muscles 140

40. Corvus corax sinuatus, outer aspect of forearm and hand,

showing relative position, origin, and insertion of the
deep layer of muscles 143

41. Corvus corax sinuatus, inner aspect of forearm and hand,

with the superficial layer of muscles dissected away,
showing the origins and insertions of the deep layer . 145

42. Corvus corax sinuatus, anterior aspect of the bones of the

left hand, showing origin and insertions of muscles . . 152

43. Corvus corax sinuatus, ulnar aspect of the bones of the

left hand, showing origin and insertions of muscles . . 152

LIST OF ILLUSTEATIONS. ^vii

FIG. lAOE

44. Corvus corax sinuatios, posterior aspect of the bones of tlio

left hand, showing where muscles are attached .... 152

45. Corvus corax sinuatus, anterior aspect of left femur, show-

ing whei-e the muscles are attached to it 167

46. Corvus corax sinuatus, posterior view of left femur, show-

ing points where muscles arise 167

47. Corvus corax sinuatus, inner aspect of left femur, showing

areas where muscles are attached 167

48. Coriithaix erythrolopha, dissection of its thigh muscles

(after W. A. Forbes) 173

49. Corvus corax sinuatus, skeleton of left posterior extremity,

showing muscular areas of attachment 178

50. Corvus corax sinuatus, head of its tibia (enlarged), to show

points of origin of muscles 178

51. Corvus corax sinuatus, left tibia and fibula, showing mus-

cular attachments 187

52. Corvus corax sinuatus, anterior view of left leg bones,

showing where their muscles are attached 187

53. Corvus corax sinuatus, left leg bones seen from behind,

showing muscular attachments 187

53 his. Ptight thtee-quartering view of the pelvis of a speci-
men of the Piiion Jay {Gyanoceplialus cyanocephalus) ,
showing the origin of the obturator internus miiscle of
the right side, o.i. Enlarged 192

54. Corvus corax sinuatus, visceral aspect of pelvis and tail

vertebrae, showing the obturator muscle and its origin . 196

55. Corvus corax sinuatus, anterior view of tarso-metatarsus,

with muscular areas shown upon it I'Ol

56. Corvus corax sinuatus, posterior view of tarso-metatarsus,

showing areas of muscular attachment 201

h

xviii LIST OF ILLUSTRATIONS.

I'-IO. PAGE

57. Corviis corax shivatuf<, summit, of tai-KO-metatarsn«, seen

fi'om above 201

58. Corvus corax sinuatus, basal joint, of hallux, seen upon

superior view 201

59. Corvus corax sinuatus, basal joint of hallux, seen from

beneath 201

60. Corvus corax sinuatus, skeleton of left foot, seen from be-

hind, showing the areas and points of attachment of
muscles 206

61. Corvus corax siiivafus, outer aspect of the skeleton of left

pelvic limb, showing superficial muscles of thigh and leg 207

62. Corvus corax sinuatus, skeleton of left foot, designed to

show where the tendons pass 210

62 bis. Outer aspect of the right pelvic limb of Geococcyx

calif ornianus 213

63. Coi'vus corax sinuatus, outer aspect of left pelvic limb,

showing second layer of muscles . , 216

63 his. Pelvic limb, right side, of (Jeococcijx californianus ; the

superficial layer of muscles removed 221

64. Corvus corax sinuatus, outer aspect of left pelvic limb,

showing third layer of muscles of thigh and leg . . . 224

64 bis. Pelvic limb, right side, of Geococcyx cali/orniccnus,

showing muscles of the third layer, and the amhiens . 231

65. Corvus corax sinuatus, outer aspect of left pelvic limb,

showing the fourth or deepest layer of muscles of thigh
and leg 236

65 bis. Pelvic limb, right side, Geococcyx californianus, show-

ing the deepest layer of muscles 237

LIST OF ILLUSTRATIONS. xix

FIG. PAGE

65 ter. Right foot of MegalcGina asiatica, showing an arrange-
ment of the plantar tendons 239

65 quat. Left foot of Gallus bankiva 239

65 quin. Right foot of Afteryx 239

66. Corvus corax sinuattis, dorsal aspect of the skeleton of the

trunk, designed to show the areas where muscles are
attached to it 260

67. Corvus corax sinuatus, cervical vertebra^ showing where

muscles are attached to them 269

68. Corvus corax sinuatus, cervical vertebrae, showing where

muscles are attached upon their lower aspects . . . . 269

69. Corvus corax sinuatus, side view of cervical vertebra^,

showing points of origin and insertion of muscles . . 269

70. Corvus corax sinuatus, dissection of muscles of head and

neck, seen on lateral aspect 274

71. Corvus corax sinuatus, six leading cervical vertebra} (en-

larged), side view, showing deep-seated muscles . . . 280

72. Corvtis corax simiatus, dorsal view of sixth, seventh, and

eighth cervical vertebrae, showing the deep muscles that

are attached to them 280

73. Corvus corax sinuatus, side view of trunk, showing dissec-

tion of superficial muscles attached to it 293

74. Corvus corax sinuatus, side view of the trunk, showing the

deep layer of muscles 300

75. Corvus corax sinuatus, muscles of the diaphragm . . . . 313

76. Corvus corax sinuatus, visceral aspect of pelvis and

skeleton of the tail, showing the muscles attached to
them 317

THE

MYOLOGY OF THE RAVEN

{Coi^viis coixix svnuatus).

INTRODUCTION.

In all highly specialized birds, and more particularly
in those possessing great power of flight, such as the
Oscines, the Haptoi'es, and many of the Anseres, the
various muscles are chiefly noted for their density,
their dark carmine colour, and the distinctness between
the fleshy portion and the tendinous. These char-
acteristics are best seen in those groups of muscles which
are brought into play during flight. As a rule, through-
out the system, the tendons are very firm, dense, and of
a bright pearly white colour ; and these, in the limbs
especially, show a marked tendency to ossify.

Having obtained a fine adult Eaven (or Crow),
showing but few or slight injuries incident to its cap-
ture, and having comjdeteli/ and carefully plucked
the specimen, it is to be immersed in 80 per cent,
alcohol for at least forty-eight hours. The student
then must supply himself with a large delf platter, for
a dissecting tray ; a grooved director, a set of hooks
and chain, a pair of wide-aperture spring forceps, and
finally, three or four suitable scalpels of various sizes.
Everything being in readiness, the first object of

2 THE MYOLOGY OF THE RAVEN.

our dissection will be to demoDstrate and expose the
cutaneous system of muscles, or the dermal inuscles.

The dermal muscles are principally of two kinds :
viz. the dermo-osseoiis, and the true de7'mals. The first-
named have their origin on some part of the skeleton,
and their insertion in the integuments ; while the second
kind both arise and are inserted in the skin or some
of its appendages.

We never find all the known dermal muscles in any
one form ; they vary with certain characteristics of the
bird. The Peacock has a special dermal muscle to act
upon the gorgeous fan which it spreads, and is com-
posed of the feathers of the lower part of the back.

Other birds having large crests, and peculiar skin
developments about the head, which require special
movements, possess the appropriate dermal muscles to
operate upon them. Again, all birds possess an enormous
system of minute muscles divided up into an infinite
number of fasciculi, to harmoniously act upon the
feather-quills, and collectively agitate the plumage.
These are quite conspicuous in the Raven, but will not
be especially dwelt upon. With an ordinary lens their
action may be studied with comparative ease in one of
the large quill-butts of the pinion, or tail.

In the Raven the apteria are entirely devoid of
muscles, and even the other dermal muscles often
appear to turn out of their usual direction to avoid
these tracts.

The muscles may be arranged, according to the usual
divisions of the body, into those of the cutaneous
system ; those of the head and neck ; those of the
trunk ; those of the upper extremity ; those of the
lower extremity ; and those devoted to the organs of
special sense.

I. THE DERMAL MUSCLES.

I. TRUE DEKMAL MUSCLES. 2. DERMO-OSSEOUS MUSCLES.

1. The dermo-frontalis. 2. The circumconcha.

4. The dermo-dorsalis. 3. The dermo-temporalis.

6. The dermo-tensor patagii. 5. The platysma myoides.

12. The dermo-humeralis. 7. The dermo-cleido dorsalis.

13. The dermo-iDectoralis. 8. The cleido-trachealis.

9. The dermo-spinalis.

10. The dermo-iliacvis.

11. The dermo-iilnaris.

1. The derm,o- frontalis} — Carry an incision through
the skin down to the bone, on the superior aspect of the
head, parallel and close to the base of the upper mandi-
ble, extending completely across. Next, from the outer

^ Selenka, in his classical work devoted to the anatomy of birds
(Bronn's Klassen und Ordnungen des Thier-Reichs, Sechster Band.
IV. Abth., Vogel : Aves. 3 u. 4 Lieferung et seq.), touches but lightly
upon the dermal system of muscles in the Class ; while even those
subsequent writers, Fiirbringer and Gadow, in their well-nigh ex-
haustive memoirs in the same field, have by no means given us a
complete history of this part of the myological system in Aves
(Bronn's Klassen, vi. Band, 7-22 Lipferung). Such being the case, I
find but few authors with whom I can compare notes, bibliographical
or otherwise, upon the thirteen muscles described above as considered
by me as belonging to the dermal system in Corvus. And I submit
them for the most pai't as they were found to exist in the Raven,

B 2

4 THE MYOLOGY OF THE RAVEN.

end of this make an incision backwards down to tlie
skull, and posteriorly to well expose the muscles in the
upper part of the neck. This straight cut should pass
about half a centimetre to the inner side of the upper
eyelid of the same side. Keflect the flap of integu-
ment thus formed, covering the top of the skull, and
carefully examine the under side of it in the median
line, where it overlies the frontal region. In old male
Kavens, I have here found a true dermal muscle, some
3 centimetres long, and a few millimetres wdde, closely
attached to the skin. So fjir, it has not been observed
by me in female birds. By its contraction, the median
feathers on the top of the head are made to lie very
flat ; at the same time those immediately above either
eye are elevated, thus giving rise to lateral crests, which
are quite well marked. In a captive Raven, an old adult
male, I have seen the bird during certain moods make
these crests stand up so as to be very evident, not to
say conspicuous.

2. TJie circumconcha.— Thin is a dermo-osseous muscle
which surrounds the periphery of the ear-conch. To
expose it, join the longitudinal incision we have already
m.ade, and carry the scalpel just through the integument
completely around the ear, a few millimetres without its
thickened margin ; — then dissect carefully up from all

simply inviting attention to the fact that my dermo-te7nporalis
appears to be the second division of the m. cucullaris of Gadow ;
the dermo-dorsalis being also a pai-t of the same ; while the
platysma myoides of my list is the first division of the m. cucullaris
of the same writer ; and finally, my dermo-tensor j^atayii is the paus
PROPATAGiALis of the M. CUCULLARIS of Furbringer. The skin
muscles in the neck of birds have been quite extensively divided up
by Gadow and Fiirbringer, described under a superficial and deep
set, and in the main as constituting various parts of their m.
cucullaris {^vonn's Klassen, vi. Band, 11 u, \'2 Lieferung, p. 214).

THE DERMAL MUSCLES. 5

sides towards it. As we detach the thickened portion
just referred to, it will be found that it contains a dermal
circular muscle, closely attached to the skin, but arising
in an evident bundle of muscular fibres which have their
origin at the outer terminus of the supraoccipital crest
(Fig. 7, cc). The ear-conch is further supjDorted by
two ligaments, attached at anterior and superior points.
3. The dermo-temporalis. — Extend the longitudinal
incision down the back of the neck to a point between
the clavicular heads, carrying it just through the skin, and
a few millimetres to the left of the median line. Care-
fully reflect back the entire flaj^, removing the skin from
the throat and the anterior portion of the chest ; and
the alar and parapatagial duplicatures of the skin are also
to be laid open. A number of dermal muscles are now
exposed. One of the most important of these is the
dermo-temporalis. On either side it is found to arise
by a broad tendinous attachment from a small depres-
sion just above and anterior to the temporal fossa. The
lower end of this attachmcDt extends down over the
sphenotic process, being but slightly wdder than it above.
From this origin the fibres of the muscle pass directly
backwards as a rather broad flat bundle, and make some
slight tendinous attachments with the temporal, which it
covers. Once clear of the skull, this muscle, as a rather
broad and thinnish band of delicate fibres, and intimately
connected with the skin, passes directly down the side of
the neck, being separated dorsad by quite an interval
from the fellow of the opposite side, while anteriorly its
margin tends to blend with the fibres of the cleido-
trachealis ; indeed a few of its fibres may run in to join
this muscular fasciculus. Opposite the shoulder-joint in
front the lower ends of the muscular fibres of the dermo-
temporalis are lost upon the skin, or blend with the

6 THE MYOLOGY OF THE RAVEN.

fibres of the dermo-tensor patagii (to be described further
along), or else, as I say, a few of them seem to merge
with the cleido-trachealis. This muscle acts as a tensor
of the lateral cervical integuments, and through its
connection with the dermo-tensor patagii, as an auxiliary
to the tensor patagii longus.

From such information as I have at hand at the
present writing, I would say that the muscle I here
describe corresponds to the temporo-alaris of Viabane, or
at least to the upper j)ortion of the temporo-alaris of that
writer. It has also received other names at the hands
of anatomists, as may be seen by consulting their works,
or such as deal with the myology of birds.

4. The dermo-dorsalis. — I give the long muscular
strip that we find in the Eaven, in the median line,
running down the neck and back, this name. It is most
prominent in the middle of the neck, where it is inclosed
in a fold of the integument. It gradually becomes lost
as we approach the occiput, as it in like manner spreads
out and disappears over the caudal region. Much fat
may overlie it, which must be carefully dissected away to
bring the muscle into view.

By its contraction the feathers along tlic median line
of the dorsum and neck are raised, which action is
assisted by the preceding muscle.

5. The plcUysma myoidcs. — This muscle is fairly well
developed in the Raven, but requires careful dissection
to bring it into view. It arises on either side, from the
lower margin of the ramus of the jaw, below the ramal
vacuity. From these points it spreads out as a very
thin, fan-like sheath, closely attached to the skin, to
meet iu a faint raphe in the median line. It will be seen
that the common muscular layer thus formed, materially
assists in supporting the lingual apparatus and the upper

THE DERMAL MUSCLES. 7

larynx, which are the chief structures found immediately
above it. When this muscle contracts, these parts will
be compressed and elevated.

6. Tlie dermo- tensor jmtugii. — That duplicature of
the common integument, of a triangular form, which is
found in the Eaven, as in other birds, between the root
of the neck and the top of the shoulder, may be designated
as the parapatagimn, in contradistinction to the pro-
patagium, or the alar integumental duplicature.

Lying within the free marginal fold of the parapatagial

drcrmio •Le,Tn,p^raZfCs

Fig. 1. — Eight lateral view of the skull and mandible ot Corvus corax sinuatus,
designed to show the origin and insei'tion of the muscles of the head,
&c. Life-size, by the author, cc, origin of the circnmconcha.

duplicature in the Raven, we discover a well-developed
bundle of muscular fibres, the mesial ends of which
spread out, to either become, as a few of them do,
inserted into the skin upon the antero-aspcct of the root
of the neck, or else pass obliquely upwards to blend with
the fibres of the dermo-temporalis. This muscle I
designate as the dermo-tensor patagii, and externally its

g THE MYOLOGY OF THE RAVEN.

fibres converge, and opposite the outer carneons extremity
of the tensor patagii longus they become converted into
a small and delicate, thongh strong tendon, which soon
thereafter blends with the tendon of the tensor patagii
longus, and in its action plays the part of an auxiliary
to it. The dermo-tensor patagii I have already alluded to
in various publications, and find in my dissection of birds
generally that it is quite a variable muscle ; and were
these variations all known and appreciated, I am confident
that they would be characters of considerable taxonomic
value. This muscle seems to correspond to the " pars
propatagialis musculi cucullaris " of Flirbringer and
Gadow, and has by several writers been described as
occurring in the species representing a variety of orders
of birds.

7, The dernio-cleido dorsalis (Fig. 8). — A beautiful
pair of muscles exist in the Eaven upon which I have
bestowed this name. Either one arises from the upper
and mesial aspect of a clavicular limb, to the extent
shown in the drawing. As the muscle passes to the
integument, it becomes closely attached to it, is directed
over the shoulder-joint and scapular region in front,
as three distinct fasciculi, diverging in a fan-like form.
The extremities may meet in the median line of the
dorsum to merge with the dermo-dorsalis.

It is very evident that the contraction of these muscles
will brace the skin over the fore-part of the back, while
if, on the other hand, they act from their integumental
attachment, they may aid to a limited extent in the act
of inspiration.^

^ These muscles seem to fulfil, in part, the function of the dermo-
transversalis of Owen, which I fail to discover in the Eaven. This
eminent authority describes this muscle for the Ajiteryx in the
following words : — " The skin covering the dorsal aspect of the lower

THE DERMAL MUSCLES. 9

8, The cleido-tracheales.- — These muscles are highly
developed in the Raven, and seem to take the place of
the sterno-'maxillaris of Owen.^ Each arises from a

two-thirds of the neck, besides being acted upon by the constrictor
colli, is braced down by a thin stratum of oblique and somewhat
scattered fibres, dermo-transversalis, which take their origin by
fascife attached to the inferior transverse processes of the sixth to
the twelfth cervical vertebrjB inclusive ; the fibres pass obliquely
upward and backward, and are insei-ted by a thin fascia into the
median line of the skin covering the back of the neck " i^Anat. of
Verts., vol. ii., p. 111).

The dermal system of muscles is highly developed in the Apteryx,
and this may account for the absence of the sterno-ccrvicalis in such
a form as the Raven, or its occurrence in a very rudimentary con-
dition, which I failed to discover. The same author says : — " The
sterno-cervicalis arises fleshy from the posterior incurved angular
process of the sternum, from the ensifoi'm pi-olongation and middle
line of the outer and posterior surface of the same bone. The
fibres pass forward, and, diverging in gently-curved lines, ascend
upon the sides of the broad base of the neck, and are inserted by a
thin but strong fascia into the median line of the dorsal integument.
This muscle is a line in thickness at its origin, but becomes thinner
as it expands ; the anterior part is covered by the posterior fibres of
the constrictor colli" {loc. cit., p. 110).

Since writing this footnote I ascertain that the sterno-cervicalis
of Owen is considered by Gadow to represent the deep layer of
his m. cucullaris (Bronn's Klassen des Tlder Reiclts, vi. Band,
p. 214).

^ " The sterno-maxillaris [in the A2)teryx'\ appears at first view to
be the anterior continuation of the preceding \sterno-cervicalis\ but
is sufficiently distinct to merit a separate description and name. It
arises fleshy from the anterior part of the middle line of the sternum,
passes directly forward along the under or anterior part of the
neck, expanding as it proceeds, and gradually separates into two
thin symmetrical fasciculi, which are insensibly lost in the integu-
ment covering the throat and angle of the jaw. It adheres pretty
closely to the central surface of the constrictor colli, along which it
passes to its insertion. It retracts the fore-part of the skin of the
neck, and also the head. Each lateral portion acting alone would
incline the head to its own side ; the whole muscle in action would

10 THE MYOLOGY OF THE RAVEN.

small circumscribed area situate upon tlie antero-inner
aspect of the middle of either limb of the furcula as a
rounded, though not large, bundle of muscular fibres.
They take their course directly uj) the neck, separated
by quite an interval. About the middle of their path,
they each flatten out and become intimately attached to
the skin and the dermo-temporalis muscle. Further
along, they become gradually narrower again, and are
finally inserted, touching each other by their inner
borders, on the anterior aspect of the superior larynx,
the trachea, and the skin over these parts. It is only
for about their middle thirds that they may be con-
sidered as true dermal muscles, and thus account for
their appearance in the present category. By their
acting in common, or each in turn, movements of
the parts would result similar to those described by
Professor Owen for the sterno-maxillwris (see footnote,
anted).

The longitudinal incision which we made down the
back of the neck must now be extended, passing only
through the skin, to the distal tip of the pygostyle.

bend the neck ; but the movements of the head and neck are more
adequately and immediately provided for by the appropriate deeper-
seated muscles, and the immediate office of the present muscle is
obviously connected with the skin. Nevertheless, in so far as this
muscle acts upon the head, it produces the same movements as the
sterno-mastoideus in Mammalia" (loc. cit., p. 111).

Since writing the footnote just quoted from Owen, 1 have ascer-
tained (Bronn's Klassen des Thier-Reichs, vi. Band, pp. 214, 215)
that Professor Gadow considers the constrictor colli of Owen to be
the superficial layer of the muscle called the cucullaris by Professor
Fiirbringer and adopted by himself, while the dermo-spincdis of
Owen constitutes the third division of the same muscle, or the m.
cucullaris, pars propatagialis, of Fiirbringer, which, as I have above
remarked, is my derma-tensor paiagii. — R. W. S.

THE DERMAL MUSCLES.

11

Reflect back on either side the integumental flaps, as
far as the sides and the Jimbs. Several muscles of the
cutaneous system are now exposed in this region.

9. The der mo- spinalis. — This delicate muscle is but

Fig. 2. — Sujierior aspect of the skull of Raven, adult $, life-.size. ]\Iandible
removed. Shows the upper edge of the insertion of the temporal muscle,
and that part of the masseter which arises on the zygoma.

feebly developed in the Raven, but a little staining in
old males will usually bring its fibres into view. It
arises in an attenuated fascia from the crests of the

12 THE MYOLOGY OF THE RAVEN.

neural spines of the first, second, and third dorsal
vertebrae, the fibres pass directly outwards, closely
attached to the skin, in rather a broad pale stratum, to
be lost over the scapular region on either side of the
body.

10. The dermo-iliacus is a parial muscle, either one
of which arises fleshy from the inner angle of the emar-
ginated portion of the antero-dorsal border of the ilium.
The fibres pass directly forwards, as a narrow ribbon, to
spread out as a thin integumental layer, to be lost
over the region of the shoulder-blade. These last two
muscles acting from their points of origin would so pull
the skin as to cause the feathers attached to it to lie
flat, or closer to the body, over the parts where they are
severally distributed. In this way they antagonize some
of the muscles described above, as, for example, the
dermo-dorsalis.

11. The dermo-uhiaris^ is exposed by still further
reflecting the skin. It is a very prominent muscle, and

1 This muscle is the pars metapatagtalis of the m. serratus
suPERFiciALis s. THORACi-scAPULARis of Furbringer, and also of
Gadow (Bronn's Klassen, vi. Band, p. 221), and represents the
tensor memhrance posterioris aloe of Wiedemann and Tiedemann,
of Kiidinger, of Selenka, and of Carlsson, while Schtipss described
it in part as the m. plicce alaris posterioris. Of it Gadow has said : —
" III. Pars meiajxttagialis. Dieser Theil ist nach Flirbringer als
aberrirende zur Haut gehende Lage des Serrat. post, aufzufassen.
Er entspringt mit zwei oder drei, selten mit nur einer Zacke
zusammen mit den Biindeln der Pars post, serrati von den Rippen,
heftet sich an die Hautduplicatur zwischen Oberarm und Rumpf
und verliert sich an den oberen Schwungfedern zweiter Ordnung.
Der Muskel is demnach ein Theil des weiter unten als M. meta-
2Kitagialis zusammengefassten Flughautspanners. Er findet sich
bei Gallus, Anser, Cygnus, Fulica, Otis, Psittacus, Raptores, Cor-
vidse, U.S.W., fehlt aber den Spheniscidse, Tubinares, Ratitse " (Bronn's
Klassen, vi. Band, p. 224).

THE DERMAL MUSCLES. 13

one of the most conspicuous of the cutaneous system.
It arises by a thin fascia from the outer aspects of the
third and fourth true vertebral ribs, at the bases of their
epipleural appendages, and from the fascia between
them. From this point it passes forwards and upwards
as a thick muscuhir bundle, loosely attached to the skin,
until it arrives in a point at the inner end of the integu-
mental duplicature behind the humerus. Here it be-
comes a small and slender tendon lying between these
skin flaps in the line of their posterior margin, as far as
the elbow-joint, where the tendon spreads out to become
attached to the fascia in the locality, and covering the
olecranon of the ulna. By extending the elbow-joint it
assists in elevating the wing.^

12. The dermo-humeralis ^ arises as a broad but

1 Owen describes for the Apteryx a dermo-costalis which " arises
fleshy, from the costal appendages of the seventh and eighth ribs. The
fibres pass forward and join those of the preceding muscle \dermo-
iliacus] to be inserted into the scapular integument " (Anat. of Vei-ts.,
vol. ii., pp. Ill, 112). I made several special dissections for the
very purpose of finding this muscle in the Raven, but my investiga-
tions lead me to believe that it does not exist in our present subject.

2 FUrbringer and Gadow have divided the M. pectoralis into
three parts, and the dermo-hume7'alis as here described for the
Raven corresponds to the third part, or the pars abdominalis
(Flirbringer).

Gadow presents us with the synonymy of the dermo-humeralis as
follows : —

" III. Pars abdominalis (FUrbringer).
Suhcutaneus abdominalis. Wiedemann, Tiedemann, § 100, No. 5.

„ „ Watson, p. 55.

Suhcutaneus thoracis. Tiedemann, § 100, No. 4. Prechtl, § 71.
Ohne Namen erwahnt. Schopss, p. 112.
Dermo-humeralis. Owen, p. 24.
Panniculus carnosus (second portion). Reid, p. 139.
Muscle des jmruo'es. Gervais et Alix, p. 401.
„ ,, Watson, p. 81."

14 THE MYOLOGY OF THE RAVEN.

barely ]:»erceptible fan-like muscle in the abdominal
integument. This is soon gathered, as it passes forwards
and upwards, towards the head of the corresponding
humerus, into a decided, long narrow fasciculus of fibres,
to be finally inserted by a very pretty little fan-like
expansion of thin glistening fascia into the tendon of
the pectoralis major just below its insertion. In the
Apteryx the dermo-humeralis " is inserted fleshy into
the proximal part of the humerus " (Owen). In either
case it depresses the humerus in the last-named bird by
acting directly upon the bone, while in the Raven it is
an auxiliary to the great pectoral muscle.

13. The dermo-2)ectoralis is a muscle that corre-
sponds to the dermo-dorsalis of the back, and is simply
the evident muscular stratum underlying the feather-
c[uills of the tracts of these appendages, that are found
on either side of the robust chest of this bird, running
in the longitudinal direction.

Either of these muscles disappears anteriorly over the
origin of the cleido-trachealis, while behind their ter-
minal fibres merge imperceptibly into the integument
after passing the hinder tips of the postpubic elements
of the pelvis. Acting from the anterior skin insertion,
these muscles raise the feathers of the chest, the action
being reversed by the muscle contracting from the
opposite extremity.

II. THE MUSCLES OF THE HEAD.

INCLUDING THE MUSCLES OF THE LINGUAL APPARATUS,
TRACHEA, AND THOSE OF THE EYE AND EAR.

Divide the skin completely around the entire beak, at
its base ; divide the platysma myoides close to its origin,
on either side, and allow its giilar portion to come away
with the integument as it is removed ; in like manner,
divide, on either side, close to their origins, the circum-
conchse and dermo-temporalis. Carefully reflect back-
wards the entire skin, so as to expose the neck for at
least its upper third. Open the bird's mouth to its full
extent, and pack sufficiently with soft tow, so as to
render the muscles of the tongue and trachea more
prominent.

The muscles here to be examined are the intrinsic
muscles of the head, and are in no way attached to the
vertebral column. They are the following : —

14. The temporal. 18. The pterygoideus internus.

1.5. The masseter. 19. The pterygoideus externus.

16. The biventer maxillaj. 20. The digastric.^

17. The entotympanicus.

1 This group of muscles (14-20 inclusive) are presented by Gadow
in his well-known memoir on the morphology of Aves in Bronn's
Klassen, vi. Band, pp. 318-325, where tliey are illustrated by

16 THE MYOLOGY OF THE RAVEN.

14. The temporal. — To thoroughly expose, this
muscle we must divide the strong lateral ligament of the
jaw, which is attached above the tip of the squamosal
process, and below to the outer side of the articular end
of the mandible. The dense fascia encasing the temporal
should also be largely dissected away.

The bulk of this fan-shaped muscle will be found
attached to, and occupying the entire temporal fossa.
Anterior to this, some of its more fleshy fibres arise from
the mesial side of the sphenotic process, and the adjacent
wall of the orbit. From this extensive origin the fibres
converge as they pass downwards and forwards, blend
more or less completely with the fibres of the masseter as
they come opposite to them, to finally become tendinous,
to be inserted in the coronoid process upon the superior
ramal margin of the mandible.

15. The massete^' (Figs. 1 and 7). — This muscle,
which may be considered a powerful auxiliary to the
one just described, is in the Raven divided into two
distinct portions. The greater mass arises by a broad
and thin tendon from the entire length of the bony ridge
above the auditory entrance, and the squamosal process

numerous handsome and useful figures on plates, and an extensive
synonymy is given, the latter being somewhat too extensive to
reproduce in the present connection. He divides his m. digastricus
s. DEPRESSOR MANDiBUL^ into three portions («, Aussere Portion ;
h, Mittlere Portion ; and c, Innere Portion), and separately defines
them. The temporal has also been somewhat similarly dealt with,
having been divided into four portions, with an accessory part.

The pterygoidal muscles are given under a single caption, the
MM. PTERYGOiDEi. Numerous prominent writers have been omitted
in so far as the synonymy goes, though the subject of these head-
muscles has been carefully handled by the author in question.

In my description above, I have essentially adopted (for the
present) the nomenclature of Owen {Anat. Verts., vol. ii.), though
subsequent dissections may incline me to alter or abandon it.

THE MUSCLES OF THE HEAD.

IV

which extends it. It also has a fleshy attachment to the
outer aspect of the body of the quadrate hone. As it
passes downwards and forwards beneath the zygoma, it

Fig. 3. — Under side of skull of Raven. Life-size. Shows the attachment of
the muscles found on this aspect of the skull. By the author.

blends with the fibres of the temporal, and makes its
first insertion by a tendinous slip to a small tubercle,
situated on the upper border of the jaw, behind the

0

18 THE MYOLOGY OF THE EAVEN.

coronoid process. The great bulk of this division of the
muscle, however, passes on to be inserted, fleshy, on the
outer side of the ramus of the mandible, nearly as far
forward as the horny beak, wdiile behind it is bounded
by the prominence of the articular extremity (Fig. 7).
Quite a dense fascia encases this division of the muscle.

The smaller part of the masseter arises, rather fleshy,
from the side and under border of the hinder fourth
of the zygoma. This auxiliary and spindle-shaped slip
terminates behind in a small, though strong, tendon,
which is inserted into the mandible on its upper border,
immediately in front of the articular end (Fig. 5, ymf).
The contraction of this part would diaw the jaw against
the quadrate, and, to a limited extent, close it. This last
action, however, is jiowerfully eS'ected by the united
contraction of the temporals and masseters.

Owen says : — " In the Cormorant, the osseous style,
movably articulated to the superoccipital, aff"ords to the
temporal muscles a more extensive origin. This, indeed,
is its essential use, for the muscles of the upper part of
the neck are inserted into the occipital bone, and glide
beneath the posterior or super-added fasciculi of the
temporalis."

16. The biventer maxiUce, on either side, is that
muscular mass which covers the mandibular articulation
behind. It arises above from two points, viz. the ridge
bounding the hinder part of the osseous ear, and secondly
the depression to its mesial side. The two heads almost
immediately blend as they pass downwards and forwards,
as a curved fleshy mass, to become inserted into the
posterior aspect of the articular end of the mandible,
including the angle (Figs. 4 and 7). To study this
muscle properly it should be transversely divided across
its belly at about its middle ; the two extremities may

THE MUSCLES OF THE HEAD. 19

then be easily dissected up,above, to its origin; below, to its
insertion.

The relations of this group, the openers and closers
of the jaw, to the surrounding structures, as the ear,
the dermo-temporalis muscle, and the quadrate, should
be carefully examined.

To expose the next set, divide the biventer maxillse as
already directed, say on the right side ; cut the corre-
sponding tendon of insertion of the temporal ; divide the
masseter as its origin ; and finally disarticulate the man-

p I e

biventer maxilho
rectus capitis posticus major

Fig. 4. — Posterior view of skull of Raven. Life-size. Shows the origin of the
muscles upon this aspect. After nature, by the writer.

dible, lifting the cranium so that we have a plain view of
its under side. Matters may be somewhat improved,
too, by severing the zygoma at its middle, and pulling
the extremities upwards.

17. The entotympanic is a small, spindle-shaped
muscle, which arises quite fleshy from the side of the
basisphenoid, and to a limited extent from the base
of the rostrum immediately beyond it. As it passes
backwards and downwards, it rapidly contracts to form

C 2

20 THE MYOLOGY OF THE RAVEN.

a double tendinous slip. One of these is inserted into a
little spine-like process on tlie upper side of the sliaft
of the corresponding pterygoid, close to its quadratal
articular end. The other is inserted into the quadrate
itself, close to the pterygoidal articulation, and to its
outer side.

When these muscles contract, they pull forward
the quadrates and pterygoids, which latter, in push-
ing against the palatines, tend to raise the superior
mandible.

18. The jyteri/goidevs internus forms, with the aid of
its fellow, the great muscular mass which is found at the
roof of the mouth. It has a very extensive origin. On
the upper side of the palatine the fibres arise over nearly
the entire surface, at first fleshy, to become tendinous
along the hinder margin of the bone. Similarly, on the
inferior asj)ect they fill the fossa there found. As the
fasciculi of this muscular mass pass backwards and out-
wards, they are augmented by others, which find their
origin upon the distal head and shaft of the corre-
sponding pterygoid and the sphenoidal rostrum for
some distance behind it. The fibres converge moder-
ately, to make an extensive tendinous insertion upon
the antero-internal aspect of the articular end of the
mandible. Mesially, they are carried out to the ex-
tremity of the inturned tip, where the tendon is very
strong ; while beyond they encroach slightly upon the
ramus, where the insertion is more of a fleshy character
(Figs. 5 and 6).

19. The pterygoideus externus is an exceedingly
neat and well-defined muscle. It arises from the outer
aspect of the extremity of the orbital process of the
quadrate. The fibres form a roundish bundle, and
slightly diverge as they pass forwards, downwards, and

THE MUSCLES OF THE HEAD.

21

outwards, to insert themselves upon the inner aspect of
the mandibular ramus, filling the fossa found in front
of the articular enlargement, and in which we find the
ramal vacuity. This latter foramen is closed in by the
pterygoideus externus in this situation, while outwardly
the masseter covers it up, the two muscles at this j^oint
being practically in contact.

20. The digastric arises on either side from an ellip-
tical, longitudinal space, of no great size, situated at the
base of the cranium, on the externo-lateral auffle of the

^•^'^^-z?

i'Ot.clg^

A^*^.

. tnji'^ ■

bivcnX^f

Fig. 5. — Left lateral aspect of mandible of Corvus corax sinuatus, slightly tipped
forwards to show muscular attachment on inner side. Life-size, vi', the
point of insertion of the zygomatic division of the masseter.

basitemporal (see Figs. 7 and 11). From this origin
the muscle passes down the side of the neck as a thin,
narrow band of fibres. Opposite the angle of the jaw
these begin to diverge, and spread out in a fan-like
form, to meet a corresponding expansion of the opposite
muscle in a median raphe, which extends longitudinally
over the superior larynx. They are still more definitely
inserted into the cartilaginous continuation of the second
basitemporal about its sides. The middle portion of
this extremely attenuated muscular stratum is very

22 THE MYOLOGY OF THE RAVEN.

dilEcult to thoroughly trace in the younger specimens
of Ravens, and it admits of our seeing the underly-
ing structures throuoh it. The delicate fibres of the
j^latysma myoides overlie it, while^ its anterior fasciculi
hlend with those of the mylo-hyoideus.

According to Mivart, in birds and reptiles the di-
gastric descends from the hinder part of the cranium to
the posterior end of the mandible, and in some birds is
divided into three portions (Elem. Anat., p. 313). As
we have just seen, it is still different in the Corvidw.

The chief use of this muscle would seem to be to
raise the hyoid apparatus and trachea against the pha-
rynx, after a morsel of food has passed that point, to
prevent its return to the mouth, and assist its passage
down the oesophagus.

III. THE MUSCLES OF THE TONGUE.

In this section I will speak of those muscles of the
Raven which 1 have found upon dissection to be, in

Fig. 6. — Under side of mandible of a Raven, designed to show the sites of
muscular attachment ; life-size, from the author's dissections.

some way or another, attached to the hyoidean apparatus,
and involved in its movements.

24 THE MYOLOGY OF THE RAVEN.

In Fig. 7, and otlier illustrations to this work, I
have made careful drawings of these parts, and en-
deavoured to present this group of muscles in such a
manner as they can best be seen and studied.

The following are presented for our examination : —

21. Mylo-hyoideus. 25. Sterno-hyoideus.

22. Stylo-liyoideus. 26. Depressor-glossus.

23. Genio-hyoideus. 27. Cerato-glossal.

24. Cerato-hyoideus.

21. The mylo-hyoideus^ (Figs. 5, 11, and 17). — This
muscle arises on the inner side of the mandible, just above

1 Professor Hans Gadow sees two parts to the mylo-hyoideus in
birds, his M. mylo-hyoideus anterior corresponding to my mylo-
hyoideus^ as shown by me in Fig. 11 of the present work (see
Bronn's Klassen, vi. Band, pp. 304-307).

To me, the m. mylo-hyoideus posterior of Gadow did not seem
to be sufficiently evident in the Baven to merit a separate desciip-
tion, and it is in no way so distinct a muscle as he figures it in liltea
darwini (I.e., Fig. 1, M.H.^a, p. 306).

Nevertheless, it has been vai'iously considered by other ana-
tomists, and Gadow offers us the following synonymy and remarks
thereon : —

" 103. 1. Grupe des M. mylo-hyoideus.

103a. M. mylo-hyoideus anterior.
M. genio-hyoidean. Vicq d'Azyr, 1773, p. 585, No. 1.
Mylo-liyoidean. Cavier. Duvernoy, p. 5, No. 1.

,, Gervais et Alix, p. 19.

Mylo hyoideus transversus. Tiedemann, § 90, No. 1.

Nitzsch, p. 134, No. 2.
Mylo-hyoideus ohliqutis. Kutorga, p. 20.
Khomhoideus juguli. "Wiedemann, p. 70.
Querer Unterkiefermuskel oder vorderer, oberfliichlicher Kiefer-

Zungenbeinmuskel. Meckel, p. 409, No. 2.
3fylo-glossus s. transversus mandibulce. Gurlt, p. 14.
Uylo-hyoideus. Schneider, p. 145 ; Gadow.
,, Watson, p. 136.

" Dieser Muskel erscheint nach Abtragung der Ilaut zwischen den
heiden Unterkieferiisten. Er entsjjringt von der Innenfliiche, oder

THE MUSCLES OF THE TONGUE. 25

the lower border and tlie intunied edge of the horny
sheath of the bill. The line of its origin occupies about

auch vom Eande des Unterkiefers mit ganz quer gericbteten Fasern,
die sich in der Mittellinie mit denen der andei'en Seite unter
BilduDg eines longitudinalen Hclmenstreifens vereinigen. Cle-
wohnlich hat der Muskel keine direkte Verbindung mit dem Zun-
genbein." [This is followed by an account of its variations in a
number of groups of birds.]

" 103b. M. mylo-hyoideus posterior.
Mylo-hyoidien. Vicq d'Azyr.
Serjnhyo'idien. Cuvier.

,, Diivernoy. •

Retrahens linguce. Wiedemann, p. 71.
Mylo-hyoideus obliquus. Tiedemann, § 90, No. 2.

,, ,, Kutorga, p. 20.

„ ,, Nitzsch, p. 135, No. 4.

Heber des Zungenbeins. Meckel, p. 409, No. 1.
Mylo-hyoideus. Gurlt, p. 14.

Retractor de Vhyoide ou serpi-hyo'idien. Gervais et Alix, p. 18.
Serpi-hyoideus + Stylo-hyoideus. Gadow, pp. 66, 67.
Retractor linguce. Watson, p. 135.

" Halswjirts vom Al. tnylo-hyoideus, ebenfalls subcutan gelegen
zerfiillt dieser, grosser Verschiedenheiten zeigende Muskel, bisweilen
in zwei Theile. Bei Rhea darwini entspringen beide zusammen
von der Aussenfliiche des hinteren Ende des Unterkiefers, laufen
liber die nach Aussen iibergreifende Insertion des M. pterygoideus,
werden breiter und theilen sich in eine vordere und eine hintere
Masse. Die letztere (J/, serjn-hyoideus) geht mit ziemlich trans-
versaler Richtung zur Mittellinie ; ihre tieferen Ziige befestigen sich
am Bindegewebe der Ventralfliiche des Larynx, die oberflachlichen
vereinigen sich aponeurotisch mit denen der anderen Seite und
strahlen zugleich auf den Hinterrand des If. mylo-hyoideus, sowie
halwiirts auf den M. constrictor colli aus. Die vordere Masse (J/.
stylo-hyoideus) ist schmaler, geht schriig vorwiirts und einwarts und
inserirt sich in dem von den beiden Zungenbeinhornern und dem
Zungenbeinkorper gebildeten Winkel ; der Insertionstheil wird
ventralwarts vom M. mylo-hyoideus anterior bedeckt."

After giving its variations in a number of groups of birds he
adds that " Bei Corvus kommt der gemeinsame Ursprungstheil

26 THE MYOLOGY OF THE RAVEN.

the anterior third of the entire length of the ramus,
and is carried forward to the posterior margin of the
horny covering of the beak, where the Latter overhi])s
the symphysis. Its wonderfully delicate fibres are di-
rected transversely to meet those of the muscle coming
from the opposite side. The two unite in a mid-longi-
tudinal raphe, which as it is produced backwards be-
comes faintly tendinous, and is finally inserted as an
attenuated aponeurosis into the under side of the hyoid,
between the first and second basi branchial, in the middle
line (Fig. 17).

The thin sheet of muscle thus formed spans and
stretches completely around the forepart of the inter-
ramal space, and by its contraction the tongue is lifted
upwards against the roof of the mouth.

This muscle is well developed in other classes of
animals. Professor Mivart found it very large in Meiw-
jwrna, and it varies considerably among the higher
Mammalia.

22. Jlie styJo-hyoideiis is the name used by Professor
Owen for a muscle which is quite conspicuous in the
Eaven. Its orioin and insertion are well shown in Fio-.
7, where the lingual apparatus is drawn down by the
dissecting chains, in order to put it on the stretch
(see various views of the mandible, where it is shown).

The stylo-hyoideus arises on the outer aspect of the
articular enlargement of the mandible, at about the
middle point. Its fibres form a rather long, transversely

hauptsiichlich von der iiusserlich den M. hiventer ')nandihulai
bedeckenden Fascie."

Having carefully dissected the iniylo-hyoideus out in some fifteen
specimens of the Raven, it seems but proper that I should say here
that my observations upon it agree with those of Owen, and in the
form in question I have thus far failed to detect a posterior mylo-
hyoideus (see Owen's Anat. Verts., vol. ii. p. 153).

THE MUSCLES OF THE TONGUE. 27

flattened fasciculus, which proceeds downwards and
forwards to be inserted by a delicate tendon into the
head of the cerato-branchial of the thyro-hyal of the
corresponding side on its upper aspect. When these
muscles contract in unison, they tend to raise the tongue
towards the palate, but if they act singly this member is
pulled to the side towards wdiich the contracting muscle
is directing it. We find that " in some birds it divides
into three or more portions : the posterior descends
obliquely forward, and is inserted into the tendinous
commissure of the mylo-hyoideus ; the middle portion is
inserted into the urohyal [second basibranchialj ; the
anterior fasciculus is inserted into the side of the
basihyal above the transverse hyogiossus. The actions
of these different portions vary according to their in-
sertion ; the first and second depress the apex of the
tongue by raising the urohyal, the third raises the tongue
and draws it to one side when it acts singly " (Owen).

23. ll^e genio-hyoideus is the largest and most
powerful muscle of the group we are now examining
(Figs. 5 and 7, and others). It arises from an extensive
area (two in some birds) on the inner side of the mandible
nearly as far forward as the horny jDart. The fibres
unite to form a broad band, transversely compressed,
which passes downwards and backwards until it meets
the outer side of the corresponding thyro-hyal, about its
middle ; this it envelops, by a spiral turn and a half, to
its very extremity.

It is quite evident that the contraction of this pair of
muscles will have the effect of protruding the tongue
from the mouth.

24. The cerato-hyoideus (Figs. 7 and 9). — I find the
muscle so named by Owen to have a somewhat differ-
ent insertion than that eminent anatomist gives it for

THE MUSCLES OF THE TONGUE. 29

the class, generally. Of it, lie says that " it passes from
the thyrohyal to the urohyal, and is therefore sub-
servient to the lateral movements of the tongue " (Anat.
Verts., vol. ii. p. 154). This duty it also performs in
the Kaven, where we find it arising on the under side of
the shaft of the epibranchial element of the thyro-hyal
by a delicate tendinous slip. This soon becomes muscular
as it passes forwards, forming a long slender belly, which
is closely applied to the hyoid. Opposite the first
basibranchial it is again a long tendinous slip, and thus
remains until it is finally inserted into a little bony
elevation on the under side of the anterior end of the
corresponding cerato-hyal.

Owing to the free lateral movements enjoyed by the
joint at the anterior end of the first basibranchial, it is
quite clear that either of these muscles acting singly
must pull the tongue to one side. But the joint referred
to also operates in the opposite direction, so that, united
action of the pair would tend to depress the tongue.

25. The sterno-hyoideus (Figs. 9 and l7). — Agreeing
with the majority of birds, we find in the Eaven either
sterno-hyoideus arising from the anterior surface of the
thyroid bone of the superior larynx. The muscular fibres
at the origin are quite fleshy, but as they converge and
pass forwards, they become rather more tendinous, and
are thus constituted as they are inserted into either
cerato-hyal at its base. Other fibres, which hold a more
mesial position, unite with the corresponding ones of the
muscle of the opposite side, to pass into and finally be
lost in the tissue composing the soft part of the tongue,
at its base. These latter are not well shown in Fig. 17,
as they are huddled together by the traction of the little
dissecting hook and chain, which pulls the muscle to one
side so that the depressor glossus may be seen.

30 THE MYOLOGY OF THE RAVEN.

If these muscles contrcact in unison, the hirynx being
the fixed point, they will evidently depress the tongue.
Acting singly in the same way, either one will likewise
depress the tongue, but also giv^e it a lateral deflection to
the side towards the muscle which is contractino;.

If they contract together, with the base of the tongue
as the fixed point, they will pull the larynx forwards, a
very necessary movement during the accomplishment of
the act of deglutition.

26. The depvessor-glossus is the name 1 have bestowed
upon a small but important muscle which Professor Owen
describes in the following words: "A small and short
muscle, which is single or azygos ; it passes from the
basihyal to the under part of the glossohyal ; it depresses
the tip of the tongue and elevates its base" {loc. cit.,
p. 154). Nameless though he left it, Professor Owen's
description answers very well indeed for a corresponding
muscle that we find in the Raven, and the one, as I say,
I propose to call the depressor-glossus. In this bird it
arises fleshy over the entire under surface of the first
basibranchial (basihyal of Owen), the fibres contract as
they pass forwards, and become tendinous between the
tongue bones, and as a delicate tendon so continue on to
finally find an insertion at their tips, where they meet
anteriorly (Fig. 17). When tliis muscle contracts, the
tongue must behave in a manner already described, and
quoted from the authority above.

27. The cerato-glossus is a small muscle we find on
the upper side of the hyoid, one on either cornua. Each
occupies, as an origin, the half of the surface of the first
basibranchial. After passing over the joint formed by
the head of the thyro-hyal, the fibres become more
tendinous, as they insert themselves along the upper
side of the shaft of the cerato-branchial element. They

de.Ztoid'

rafo/tiivg

tJ^eMS

Fig. 8. — Oblique view from the iBner side of the furcula, coracoid, and scapula
of the Raven, showing the areas devoted to the muscles which take
origin from these bones. Life-size.

Fig. 9. — Under view of the hyoid arches of a Raven, with the upper part of the
windpipe, in situ. Designed to show the origin and insertion of certain
muscles of the tongue, m.h., lingual attachment of the inylo-hyoideus.
Life-size.

Fig. 10. — Superior view of the bones of the upper larynx and half- rings and
rings of the trachea. Shows the origin and insertion of the two
muscles which control the rima glottidis. x 2. All the figures drawn
by the author from his own dissections.

32

THE MYOLOGY OF THE EAVEN.

Fig. 11. — ITnder view of the head of a Raven, with mandible in situ, hut with
hyoid apparatus and other parts carefully removed. Dissected to show
the muscles which occur upon this aspect ; the left side (the right in
the figure) shows the origin and insertion of the jjiarygoideus intemus.
The opposite side is dissected to show the entotympanicus and plery-
goidcus cxternus. The anterior two-thirds of the inylu-hyoidc/us is also
shown, and its mode of attachment. The digastric is cut off close at
its point of origin. This figure should be carefully compared with
Fig. 3 of the present memoir. Life-size, by the author from his own
dissections.

•

here glide beneath the thyroid bone of the superior
hirynx. Just beyond this point they become fully

THE MUSCLES OF THE TONGUE. 33

tendinous, and are finally inserted as a distinct tendon
at the muscle's termination on the shaft of the element
just alluded to, and in line with the last-named fibres.

According to Professor Owen, Dr. Salter proposed the
name of this muscle which I have here adopted for it.
The former anatomist describes it as " a short muscle,
which arises from the junction of the basihyal with the
urohyal, and is inserted into the thyrohyal " (loc. cit.,
p. 154).

Professor Owen figures it in a Fieldfare, but makes an
incorrect reference to his figure, and the muscle is shown,
apparently, as being pulled out of its normal position,
in order to show the stylo-hyoideus.

When these muscles contract, they will elevate the
long cornua of the hyoid arches, and press them up
against the skull.

In addition to the muscles already described as be-
longing to the tongue, there is still another pair, found
by Owen in a Woodpecker (shown in Fig. 12 at h),
which he has named the cerato-trachecdes, and says :
" They arise from the trachea about eight lines from
the upper larynx, twist four times spirally round the
trachea, and then pass forward to be inserted into the
base of the thyro-hyals. This is the principal retractor
of the singular tongue in this species " (Anat. Verts.,
vol. ii., p, 154).

Unfortunately, Professor Owen does not mention in his
work the species of Woodpecker in which he found this
interesting pair of muscles. He speaks of it, however^^
in such a way as to lead one to suppose that it occurs^Jr
throughout the family. Dissections made upon our
American forms, with a view of looking further into this
matter, would be well worth the time and labour of the
comparative anatomist.

D

34 THE MYOLOGY OF THE RAVEN.

Eeferrino; aaain to Professor Gadow's estimable work
in Broun's Klassen des Thier-Reichs (vi. Band, pp.
307-317), we find that lie has described and proposed
tlie following nomenclature for the muscles of the lingual
apparatus in birds. In presenting this I refer to the
plates of this writer, although his figures are not here
reproduced. It will be observed that he has bestowed
very diff"erent names upon the muscles as compared with
Owen's nomenclature, the authority at my hand when
the present volume was written.

" II. Muskeln des Zungengeriistes. (Taf. xxxii., xxxiii., nnd

Holzsclinitte.)

" 104. System des M. sterno-hyoideus.

Thyreo-hyoideus. Vicq d'Azyr ; Gervais et Alix, p. 18.

M. laryngo-hyo'idei. Tiedemann, § 90, No. 5.

Hyo-thyreoideus. Kutorga, p. 20. Gi;rlt, p. 15.

Thyreo-hyoideus. Nitzsch, p. 136, No. 5.

Thyreo-glossus. Nitzsch.

Thyrohyoid. Watsou, p. 136.

M. ypsilo-trachealis s. dejwsssor arterice aspera; snpe7[ficialis, major.

Meckel, 6 Tbeil, p. 284.
Sterno-hyoideus. Nitzsch; Giu-lt, p. 15.
Tracheo-sternalis. Gadow, Tenuirostres, p. 68.
Oino thyreoid ei. Huber.
Les cerato trachees. Cuvier.
Cerato-trachei. Tiedemann, jx 122.
Tracheo-glossi. Huber.
CeratO'trachealis. Owen .
Trachea hyo'id ten. Duvei'noy.
Tracheo-hyoideus. Gadow, p. 62.

" Ein echter, an das bei den Eeptilien allgemeine urppriingliche
Verhalten anschliessender M. sterno-hyoideus ist noch bei Ajjteryx
vorhanden. Der breite Muskel entspringt vom vorderen Rande der
Unterfliiche des Thyreoid-Knorpels des oberen Kehlkopfes und vom
ganzen Innenrande der Zungenbeinhorner. Seine Fasern sind
abwiirts gerichtet, verbinden sich theilwcise mit denen der anderen

THE MUSCLES OF THE TONGUE. 35

Seite ill der ventralen Mittellinie und bilden dann eine ziemlich
dicke und breite Muskellage, welche ventral und seitlich die Trachea
lose umgiebt, ohne jedocli an derselben befestigt zu sein. Nahe
der Brust angelangt, theilt sich die bis dahin vereiiiigte Masse
in eine rechte und eine linke Hiilfte, deren jede sich am ganzen
Seitenrande des Sternums und auch am grcissten Theile des hinteren
oder caudalen Randes des Brustbeines inserirt. Die Brust- und
Schultermuskeln werden natUrlich von den theilweise nponeuro-
tisch gewordenen, flachen Insei-tionstheilen des M. sterno-liyoideus
bedeckt.

" Der Muskel wird durch Zweige des N. hypoglossus, und weiter
abwiirts von den meisten Cervicalnerven aus innervirt.

" Als eine mediane, innere, von dem eben beschriebenen Muskel
abgetrennte Masse ist ein Muskel aufzufassen der bei Apteryx mit
geringer Ausdehnung fleischig vom Coracoid, nahe dessen Yerbin-
dung mit dem Sternum entspringt und sich streng median- und
kopfwarts gei-ichtet, etwas oberhalb der Theilung der Trachea in
die Bronchien an der Trachea befestigt ; von dort begleitet er die
Trachea, an derselben seitlich besfestigt, wird im Bereich des oberen
Drittels der Luftrcihre sehr schwach, schwillt aber nahe dem
Thyreoidknorpel wieder an und inserirt sich an dessen Seiten- und
Unterflache. Wir bezeichnen ihn als 31. sterno s. coracothyreoideus .
Dieser Muskel wird ausschliesslich durch einen Ast des N. hypoglossus
innervirt, der bis in die Brust hinabsteigend in dem Muskel
verfolgbar ist.

" Andere das Sternum und den Schulterglirtel mit der Trachea
oder mit dem Zungengeriist verbindende Muskeln besitzt AjMryx
nicht. Das bei diesen Ratiten bestehende Yerhiiltuiss giebt uns
Aufschluss Uber die complicirteren Zustande bei den andern Vogeln.
Aus den beiden Muskeln des Aj^teryx sind mehrere entstanden, von
denen aber nur die mit dem Zungengeriist in Verbindung stehenden
hier besprochen werden, wahrend die iibrigen, als in den Dienst der
Respiration und Stimmbildung getreten, bei jenen Organen er-
schopfend beliandelt werden. Ein M. sterno-liyoideus findet sich
ausser bei Apteryx bei manchen Carinaten. Bei Prosthemadera
wird er jederseits durch ein schmales Band gebildet, welches im
Winkel der Symphyse der Furcula entspringt tind direkt, ohne andre
Verbindungen einzugehen sich an der Basis des Zungenbeins und
zwar mehr an dessen Dorsalseitenrand inserirt ; nur wenige Faseru
befestigen sich an der Wurzel der Zungenhorner.

" Bei Meleagris ist nur die bei ApMryx als innere erwahnte mediane

D 2

36 THE MYOLOGY OF THE RAVEN.

Portion voihanden. Diesel be kommt vom Pi'oc. lateralis anterior
des Brustbeins, geht zur Seite der Trachea, wird selir schwacli und
schwillt am obern Ende wieder an, um sich am Larynx und am Os
urohyale zu befestigen ; ausserdem aber ist sie nahe dem Larynx an
jeden der TrachealriDge befestigt, oder eigentlich kommt von diesen
Ringen und geht zum Urohyal, den benachbarten Theilen des
Basihyal und der Zungenhorner. Als eine deatliche Abspaltung des
angen Muskels besitzt Meleayris noch einen, der sich vom ventralen
und seitlichen Rande des Thyreoidknorpels und vom Seitenrande
des ersten Trachealringes selbstandig und frei an der Seite des
Halses her abzieht, in der Nahe der Schulter aber ventral wiirts mit
dem der anderen Seite convergirt und eine sehr dlinne, ziemlich
schlUpfrige Schicht von Bindegewebe ohne Muskelfasern bildet, die
sich allmiilich an der Ventralflache des untern Endes der Luftrlihre
anheftet. Diese beiden langen Muskeln von Meleagris werdea von
einem Zweige den N. Ityiwjlossus begleitct.

" Denkt man sich nun, wie das iibrigens schon bei Afteryx und
Meleagris angedeutet ist, dass die Fasern des M. sterno hyoideus und
claviculo-hyoideus im Bereich des mittleren Halsdrittels verkiim-
mern, so zerfallen ebengenannte Muskeln je in einen obern Theil,
der dann als tracheoJtyoideus und traclieo-laryvgeiis superior und in
einen untern Theil, der als sterno- s. coraco- s. cleido-tracJiealis und
tracheo-lanjngeus inferior betrieben werdeu kann. Jeder derselben
kann wieder in Unterabtheilungen zerfallen.

"So haben wir bei Rhea folgende Muskeln : L Ein M. tracheo-
aryngeus superior. Derselbe erstreckt sich von der Yentralfliiche
des Hinterrandes des Thyreoids auf die Seitenfljiche des obern Drittels
der Trachea. 2. Sein vorderster Theil ist ein Thyreo-glossus s.
hyoideus, denn er verbindet die Yentralflache des Thyreoids mit dem
Hinterrande des Os basihyale. 3. Von der Seitenmuskulatur der
Trachea lust sich ein wohlentwickelter bandartiger Muskel ab,
der sich am obern Ende des ersten Gliedes des Zungenbeinhornes
befestigt, daher M. tracheo -hyoideus zu nennen ist. 4. Die Brust-
partie derselben Muskelmasse ist sehr diinn geworden und verliert
sich in der Haut des Halses, ohne mehr die Brust zu erreichen.
Ausserdem ist 5, ein Sterno-tracltealis vorhanden und ein daraus
difterenzirtes Paar Syrinx-Muskeln.

"Bei Nycticorax griseus besteht ein dem 31. tracheo hyoideus
und thyreo-hyoideus von Bhea vergleichbares Muskelpaar, das mit
einigen Ziigen von der Trachea, hauptsiichlich aber vom Thyreoid-
Knorpel zum Basihyal geht. Die Luftrohre wird nicht von Mus-

THE MUSCLES OF THE TONGUE. 37

keln begleitet, die betreffenden Theile sind daher ansgefallen, und
nur ein M. sterno-trachealis nebst einem primitiven Syrinxmuskel
oder M. tracheo-hroncldalis ist vorhanden.

" Bei Cacatua roseicajnllus konimt ein diinnes Muskelband vom
Larynx und vom Zungengerlist ; ein Theil begleitet die Luftrohi'e
bis zu den Syrinx- nnd Tracheo-clavicular-Muskeln und wird wie
diese nur vom N. hypoglossus innervirt ; ein dickerer, mehr seitlicher
Theil breitet sich bald auf der Halshaut aus, verwebt sich dort mit
den Hautmuskeln und erreicht das Brustbein oder den Schulter-
gurtel nicht. Hauptsiiclilich von einem langen Zweige des N.
hyjwglossus versorgt, ei'halt er weiter unten auch Zweige aus den
Cervicalnerven,

" Beim Griinspecht (Taf. xxxiii., Fig. 35a) [not here I'eproduced],
kommt jederseits ein diinner Muskel vom Schultergurtel, begleitet
die Trachea in ihrer ganzeu Lange und stosst erst nahe dem Kehl-
kopfe mit dem der anderen Seite zusammen, worauf sich beide am
Thyreoid und dem medianen Zungengerlist inseriren. — Ein zweiter
Muskel kommt von der Clavicula und geht etwas seitlich von dem
vorigen gerade kopfwarts als ebenfalls schmales Band und inserirt
sich am Thyreoidknorpel und am Grunde des Zungenbeinhornes
seiner Seite.

" Ein dem M. tracheo-hyoideus entsprechender Muskel erreicht bei
den Spechten seine hijchste Entwicklung. Er entspringt jederseits
von der Trachea unterhalb des Kehlkopfes, windet sich dann
mehrere Male (beim Griinspecht viermal) lose um die Luftrbhre
und geht dann an die Basis des Zungenbeinhornes. Bei weit
herausgestreckter Zunge entrollt sich dieser eigenthiimliche Miiskel
seine Funktien ist Zurriickziehen der Zunge.

" Wir konnen daher diejenigen Zungenmuskeln, welche entweder
mit dem Brustbein und Schultergurtel, oder mit der Luftrijhre und
dem oberen Kehlkopfe zusammenhangen, folgendermassen zusammen-
fassen.

" I. Gruppe.

" Vom Sternum zum ganzen Zungenhorn, und zum Basihyal :
Sterno-hyoideus ; Afteryx.

" Von der Clavicula hauptsachlich zum Zungenkorper ; Cleido-
hyoidei ; Prosthemadera, Ptilotis, Pici.

"Dieselben Muskeln erreichen aber die Brust nicht, sondern sind
an der Halshaut befestigt ; Papageien.

" II. Gruppe,
" Von der Trachea zum Zungengeviist und z"\var zvim Cerato-hyal ,;

38 THE MYOLOGY OF THE RAVEN.

Tracheo-hyoidei ; Nectarinia ; Pici ; Grallaj ; Rasores ; Eaptores :
Rhea.

" Vom Larynx (Thyreoid) und theilweise auch von der Trachea
zum Basihyal, Entoglossum nnd Urohyal ; Thyreo-hyoidei ; Anser ;
Gallus : Rasores; Ptilotis ; Rhea; Spheniscidse.

" 105. M. GENIO-IIYOIDEUS.

M. genio-liyoldes. Steno.

Le -muscle conique de I'os hyoide. Vicq de Azyr, 1773, p. 535.

Protahens linguce. Wiedemann, p. 71.

Mm. conici ossis hyoidei. Tiedemann, § 90, No. 3.

,, ,, Kutorga, p. 20.

Tiefer Vorwartzieher oder Kinnzungenbeinmviskel. Meckel, p. 409,

No. 3.
Mylo<erato-hyoidien. Duvernoy.
Genio-hyoideus. Gurlt, p. 15.

,, Nitzsch, p. 135, No. 3.

,, Gadow, Tenuirostres, Froc. Zool. Soc, 1883, p. 67-

Mylo-ceratoideus anterior et j^osferior. Nitzsch, in seiner Tafeler-

klarung, p. 151.
Genio-ceratoideus. Huber.

Trolracteiir de V hyoide {genio-hydidieti). Gervais et Alix, p. 18,
Protractor lingrioi. Watson, p. 135.

" Dieser Muskel entspringt vom inneren, nnteren oder vom oberen
Rande des TJnterkieferastes, ungefiihr in dessen Mitte und geht
bandformig gerade nach hinten, ventral von dem M. tnylo-liyoideus
anterior, aber dorsal oder in derTiefe von M. mylo-liyoideus posterior
gelegen, an die vorwjirtsschauende Seite der Zungenbeinhorner, nm
die er sich theilweise herumwindet ; ihr Enddrittel ist ganz von den
Muskelfasern umgeben. So verhalt es sich bei vielen Yogein, z.B.
bei Corvus, Anser, Procellaria, Spheniscus. Haufig jedoch (A^ec-
tarinia, Otis) zerfallt das vom Unterkieferkommende Band in zwei,
von denen das eine sich wie gewohnlich um das Zungenbeinhorn
herumwickelt, wahrend das andere sich nur an der iiusserten Spitze
desselben befestigt, beide Theile sind aber von einer gemeinsamen
schliipfrigen Scheide umgeben, wodurch ihr Zusammenwirken und
zwar ausschliesslich in der Richtung des Knochens, gesichert ist.
Bei Prosthemadera war der Ursprung des Bandes auf die Aussen-
flache des Unterkiefers geriickt.

" Bei den Papageien ist der Muskel ganz getheilt. Der vordere
(Taf. xxxii., Fig. 30d) [not re-figured here] entspringt vorn an der

THE MUSCLES OF THE TONGUE. 39

inneren Fltiche des Unterkiefers vind geht zum letzten Drittel des
ersten langen Gliedes des Zungenbeinhorns, erstreckt sicli audi wohl
etwas zum zweiten, umwickelt aber bei Papageien das Zungenbein-
horn nicht so wie bei vielen andern Yogeln. Der hintere Theil
(Fig. 30c) [not re-figured here] entspringt weit vom vorderen ent-
fernt am unteren Eande des Unterkieferastes etwa in der Mitte
seiner Lange und geht an das sehr kurze zweite Glied des Zungen-
beinhornes seiner Seite. Nitzsch schlagt fiir diese beiden Muskel-
theile die Namen M. mylo-ceratoideus anterior und posterior vor.
Bei Rhea sind ebenfalls zwei ganz getrennte Bander vorhanden.
Das vordere entspringt aus dem Kinnwinkel und stosst in der Mit-
tellinie mit dem der anderen Seite zusammen ; es inserirt sich am
Ende des ersten Gleides des Zungenbeinhornes. Das aussere Band
ist schmiiler und wickelt sich um die iiusserste H'alfte des letzten
Gliedes des Zungfenbeinhornes.

"106. M. GENIO-GLOSSUS.

M. myloglosse. Duvernoy, p. 6, No. 5.
M. genioglosstis. Nitzsch.

" Dieses Muskelpaar scheinen nur wenige Vogel zu besitzen. Bei
den Papageien entspringt es vorn dicht neben der Mittellinie der
inneren Kinniiiiche, wird vom M. mylo-hyoideus bedeckt und begibt
sich als diinnes Band an den Seitenrand des hinteren Endes des Os
entoglossum. Nach Nitzsch fehlt es bei vielen anderen Vogeln, oder
es ist, wie bei den Pavibvogeln, wo es im Frenulum der Zunge
vom Rachen aus durchscheint, so schwach, dass es leicht Ubersehen
wird.

" Den Penguinen, Trappen, Gansen, Kriihen, Hlihnern, Spechten
Kolibris, Nectarinien scheint es in der That zu fehlen.

" Bei Procellaria ist der Muskel sehr diinn, liegt der Mundschleim-
heit an, und erstreckt sich vom seitlichen hinteren Theile des Os
entoglossum diinner werdend und mit dem der andern seite con-
vergirend zum Kinn."

" 107. M. OERATO-GLOSSUS.

Cerato-glosse. Cuvier.

Cerato-glosse. Duvernoy.
Cerato-glossus. Tiedemann, § 91, No. 1.
,, Kutorga, p. 21.

40 THE MYOLOGY OF THE RAVEN.

Gerato-glossus. Watson, p. 134.

„ Gadow, p. 67.

Nieder- und Seitwartszieher der Zunge. Meckel, p. 408, No. 1.
C eratoglossus inferior s. hasioglossus lateralis und superior. Nitzscli,

p. 136, 137.
Basioglossi inferiores et superiores. Kutoi-ga, p. 21.
Vhyo-glos&e. Gervais et Alix, p. 18.
Grund ZuDgenmuskel (basio-glossus). Gui-lt, p. 1.5.
Griffel- oder Zungenbein-Zungenmuskel (.s/v//o-A?/ow/ew.s). Gurlt, p. 15.

" Im allgemeinen entspringt dieser stets vorhandene Muskel
fleischig von der oberen Fliiche des ersten Abschnittes des Zungen-
beinhornes und inserirt sich mit einer langen, deutlichen Sehne am
Seitenrande des Os entoglossum. Ha,u6g zerfiillt er in zwei Theile ;
der eine kiii'zere geht bei den Hiihnern vom Zungenklirper (Os en-
toglossum) bis fast zur Spitze der Zunge, die er herabkriimmt, er
liegt dann auf der Ventralseite der Zunge ; der andere liingere
kommt vom Zungenbeinhorn und endigt sehnig a,m Grunde des
Kbrpers." [This account of the cerato-glossus (the cerato-glossal as
described by me above for the Raven) is completed by Gadow by a
few words upon its variations as found by him in a number of
groups of birds.]

" 108. M. CERATO-HYOIDEUS.

C erato-JiydicUen. C uvier.
Gerato-hyoideus. Tiedemann, § 90, No. 4.

,, Ivutorga, p. 21.

„ Meckel, p. 409, No. 4.

„ Nitzsch ; Gurlt, p. 14.

,, Duvernoy.

Ceratoidien transverse. Gervais et Alix, p. 19.
Cerato-transverse muscle. Watson, p. 134.

" Ein kurzer Muskel, der fleischig von der Innenseite des ersten
StUckes des Zunge nhornes entspringt und mit schrag vorwarts und
median gei-ichtetem Yerlauf an den luipaarigen Stiel des Zungen-
geriistes (Os urohyale) inserirt, haufig verbinden sich dabei die
gegenseitigen Muskelziige.

" Verwebung mit dem ventral gelegenen J/, uiylo-ltyo'ideus pos-
terior ist gewohnlich. — Fehlt aus Os urohyale, Avie bei A'hea und bei
Platalea, so ist auch der Muskel nicht vorhanden, letzteres gilt aber

THE MUSCLES OF THE TONGUE. 41

auch bei vielen Yogeln, welche wie der Flamingo, Storch, manche
Eaubvogel, Prosthemadera, Nectarinia den unpaarigen Stiel be-
sitzen.

" Bei Eudyptes chrysocome ist der Muskel recht lang ; er kommt
vom zweiten Drittel des Zungenbeinhornes nnd verbindet sich anf
dem Urohyal mit dem mylo-hyoideus. Er scheint eine den Vogeln
eigenthiim-Differenzirung des M. cerato-glossus zvi sein.

" 109. M. HYPOGLOSSUS.

" Jederseits aus einem oder zwei Muskelcben bestebend, die von
der Uuterflache des Os basibyale entspringen und neben einander
auf der TJnter- nnd Seitenflache des Os entoglossum sicb inseriren.
Sie werden durch einen Zweig des Ramus lingualis N. hypoglossi
innervirt und sind hochstwahrscheinlich als tiefere, etwas selbstiindig
gewordene Tbeile des M. ceratoglossus aufzufassen.

"a. M. Hypoglossus obliquus.

Hyoglosse transverse. Cuvier ; Duvernoy.

Hyoglossus obliquus s. j^cltvus. Tiedemann, § 91, No. 2.

„ „ Gurlt, p. 15.

Heber der Zunge. Meckel, p. 408, No. 2.
Hypoglossus obliquus. Nitzsch.
Hyo'idien transverse. Gervais et Alix, p. 18.
Transverse hyoid muscle. Watson, p. 136.

" Jederseits ein kleiner Muskel, der vom Ceratoglossus bedeckt
wird ; sie entspringen nebeneinander von der untei^en Fliiche des
Zungenbeinkiirpers (Basi-hyal) mit schiefen Fasern, scblagen sich
um dessen Seitenrand und setzen sich an den hinteren Seitenrand
des dem Basi-hyal aufsitzenden Os entoglossum oder Zungenkern.
Bei Procellaria und Spheniscidce sind die beiden Muskelcben ziemlich
transversal gerichtet, mehr liinglich dagegen bei den Papageien
(Taf. xxxi., Fig. 36 und xxxii., Fig. 34) [figures not reproduced here].
Bei Rhea sind diese Muskeln mit dem sie ganz bedeckenden M.
cerato-glossus verwachsen. Bei Passerinen habe ich sie nicht be-
merkt ; bei Pelecaiius und Sula, die eine sehr verkiimmerte Zunge
haben, scheinen sie ganz zu fehlen.

*' b. M. HYPOGLOSSUS RECTUS.

Hyoglosse droit. Cuvier ; Duvernoy.

Hyoglossi interior s. rectus. Tiedemann, § 91, No. 3.

Musculi linguales hiferiores. Ivutorga., p. 21.

42 THE MYOLOGY OF THE RAVEN.

Zungenbeuger. Meckel, p. 408, No. 3.
Hypoglossus rectus. Nitzsch.

" Liegt wie der M. Jiyjwglossiis obUquus an der unteren Fliiche
der Zunge. Er entspringfc nahe der Mittellinie etwas vorwiirts vom
M. H. ohliquus und inserirt sich nahe dem Vorderende des Os
entoglossum.

" Am deutlichsten entwickelt bei Papageien, fehlt er sehr vielen
"Vogeln, und ist uberhaupt nichts weiter als die vordere Verliin-
gerung des M. hyjwglossus ohliquus. Hiermit stimmt seine
Innervation Uberein."

Note. — These are all the lingual muscles alluded to by Professor
Gadow in Bronn's Thier-Reichs, and a moment's comparison will
show the corresponding ones as I found them in Corvus, and also
such as are not found among Passerine birds. The following seem
to be the correspondences to which I allude : —

Shufeldt.

Gadow.

Mylo-hyoideus =

M.

mylo-hyoideus anterior.

8tylo-hyoideus =

M.

mylo-hyoideus posterior.

Genio-hyoideus =

M.

genio-hyoideus.

Cerato-hyoideus =

M.

cerato-hyoideus. '

Sterno-hyoideus =

M.

sterno-hyoideus.

Depressor-glossus =

M.

hypoglossus obliquus (?).

Cerato-glossus =

M.

cerato-glossus.

What I take to be the M. hyjyogloss^is ohliquus of Gadow appears
to be sufficiently evident in Corvus, and further than that genus I
have not as yet looked into the matter.

IV. THE MUSCLES OF THE AIR PASSAGES.

As in the vast majority of birds, the rings and half-
rings of the trachea of the Raven are completely ossified
in the adult individual. This applies also, with equal
truth, to the several parts of the superior larynx. These
cricoid bones and the pair of arytenoid bones are so well
known as to require no special description from me here.
Their arrangement is very well shown in Fig. 10, where
they are represented, in situ, double the size of life.

Anteriorly, the upper larynx is composed of one shield-
shaped osseous plate, the thyroid, which rests upon the
upper side of the second basibranchial, in the fork of
the greater cornua of the hyoid.

This plate is also seen in Figs. 9 and 10.

The bronchial half-rings, and those of the lower larynx,
likewise ossify, but not quite so completely as do the
tracheal rings above them.

In the Raven we have devoted to the superior larynx
two muscles —

28. The constrictor-glottidis. 29. The thyreo-arytenoideus.

And to the lower larynx seven more, viz. —

30. The tracheo-lateralis. 34. The bronchialis posticus.

31. The broncho-trachealis posticus. 35. The bronchialis anticus.

32. The broncho-trachealis anticus. 36. The sterno-trachealis

33. The broncho-trachealis brevis.

Tig. 22.

Fig. 12. — Tongue and salivary glands, "Woodpeckir. i and k, tlie glands, the

latter opened to show internal structure ; /;, the four turns of the cerato-

trachea.Ic.9 around the trachea. (After Owen.)
Fig. 13. — Front view of the lower larynx and bronchi of a Raven, g, part of the

jieripheial surface of the tAUijtanifnrni juenibrnne. The muscles above

are named in Fig. 14. (After Owen.)
Fig. 14. — Side view of the same parts that are shown in Fig. 13. a and/,

diverging fasciculi of the tradieo-latcralis muscle ; the fasciculus a is

THE MUSCLES OF THE AIR PASSAGES. 45

the hi'oncho-trachcalis posticus, and the fasciculus / is the broncho-

trachealis anticus muscle ; b is a separate muscle, the broncho-trachealis

brcvis ; c, the broricMalis posticics ; c, the bronchialis anticus ; d, the

stcr7io-tracliealis . (After Owen.)
Fig. 15. — Lower larynx of a Parrot, a, tensor loncjus (jJotticlis muscle, b, tensor

brevis rjlottidis ; c, a narrow muscle passing from the tracheal to the

bronchial half-rings. (After Owen.)
Fig. 16. — Organ of hearing, Owl. /, the tcjisor tympani muscle ; e, the Eustachian

tube ; li', the superior semicircular canal ; c, tympanum, and b is its

I^eriphery. (After Owen. )

28. Tlie constrictor glotticUs. — The inner margins of
the riiiia glottidis in the Raven are formed by the
mesial borders of the arytenoid bones. These latter
articulate posteriorly with the middle segment of the
cricoid, their anterior ends being free. Now the con-
strictor glottidis muscle arises, on either side, from the
superior and longitudinal line of the thyroid plate.
From this origin it sweeps outwards, then upwards, and
finally inwards in a gentle curve, to be inserted all along
the inner margin of the corresponding arytenoid bone,
and to the apex of the mid-cricoidal segment.

When this pair of muscles firmly contract together,
they efficiently close the rima, and draw the apices of
the arytenoids to the middle line, and close against the
upper surface of the thyroid plate. This action of the
constrictor glottidis is opposed by the next muscle, the
opener of the rimci.

29. The thyreo-arytenoideus (Figs. 10 and 18). — This
pair of muscles arise, one on either side, from the entire
outer margin of the thyroid plate, and the corresponding
margin of the outer cricoid bone (Fig. 10).

The fibres pass inwards, directly across the opening of
the windpipe, to be inserted all along the outer margin
of the arytenoid bone of the same side, and the cor-
responding border of the central cricoid piece. It is
evident that the contraction of this pair will tend to
pull the arytenoid bones outwards, and thus open the
rima glottidis. .

Fifj.lj.

Fig. 1 7. —The under side of the hyoid arches of a Raven, with the superior larynx
and upper part of the windpipe, in situ. Designed to show the attach-
ment of the muscles of the parts ; the sterm-hyoidcus and the dcido-
trachcalh are pulled outwards by dissecting hooks. Life-size, by the
author from his own dissections.

Fig. 18.— Superior view of the same parts, with lower muscles removed, but
showing those attached to the upper side of the hyoid and superior
larynx. The rima glottidis is represented as closed, with the arytenoids
in contact. The elliptical aperture, which always remains in front is
closed by the upper surface of the thyroid plate" beneath it.

THE MUSCLES OF THE AIR PASSAGES. 47

" From the simplicity of the structm'c just described,
from the situation of the superior hirynx with relation to
the rictus or gape of the bill, and from the absence of lips
by which this might be partially or entirely closed, it is
plain that it cannot be considered as influencing the voice,
otherwise than by dividing or articulating the notes after
they are formed by the lower larynx. The superior larynx
presents, indeed, but few varieties in the different species
of Birds ; and these relate chiefly to certain tubercles in
its anterior, which vary in number, and do not exist at
all in some species, as the Singing Birds ; being chiefly
present in those birds which have a rough, unmusical
voice. In the Pelican, the Gigantic Crane, and most of
the Rasoi'es, a process extends backward in the cavity
of the upper larynx from the middle of the posterior
surface of the thyroid cartilage, and seems destined to
give additional protection to the air-passages " (Owen).

To examine the lower larynx, we must carefully free
the trachea from the strong; connective-tissue which sur-
rounds it between the limbs of the furcula ; then cut
through the clavicular arch close to the hypocleidium,
through the pectoral muscles, and down through the
sternal body close to the carina. By means of this
incision carefully open the chest. Next, after studying
the relations of the beautiful pair of sterno-trachealis
muscles, nip off the costal processes of the sternum, and
free them from attached structures. Cut througii the
bronchi close to the lung, on either side, then lift out the
lower larynx for examination.

30. The traclieo -lateralis — or either one of a jDair of
muscles of that name, for all seven muscles of the lower
larynx of a Kaven are in pairs — is formed by the union
of the broncho-trachealis anticus and the broncho-
trachealis posticus, about one and a half centimetres

48 THE MYOLOGY OF THE RAVEN.

above the Infurcation of the bronchi, on the lateral
aspect of the trachea. It ascends the side of the trachea
for its entire length, as a narrow ribbon of muscle closely
attached to the rings, until it arrives near the superior
larynx, when it spreads out like a fan, and its faint and
delicate fibres attach themselves to the side of the
upper end of the windpipe. Contraction of this pair
of muscles must shorten the trachea by closely ap-
proximating the upper and lower borders of its rings.
It also gives some support to this tube by acting as a
brace to its sides.

31. The broncho -tracheaUs posticus is the hinder
fasciculus formed by the forking of the last-named
muscle. Its fibres pass downwards and backwards, and
are inserted into the end of the third half-ring of the
same side.

32. 21ic hroncho-trachealis anticus is the remainino;
limb of the bifurcation of the lower extremity of the
tracheo-lateralis. Its fibres descend downwards and
forwards to insert themselves upon the anterior extremity
of the third half-ring of the corresponding side (Fig. 19).

33. The hroncho-trachealis hrevis is, upon either side,
a strong, straight, subcylindrical bundle of muscular fibres
which arise above just beneath the broncho-trachealis
anticus muscle, descend rather obliquely across the larynx,
to become inserted into the hinder end of the second
bronchial half-ring (Fig, 14, h). This muscular slip, as is
shown in Fig. 19, is considerably larger than the broncho-
trachealis posticus, which passes down, for the major
part of its descent, alongside of it.

34. The hronchialis posticus is a chunky little spindle-
formed muscle, situated below the other muscles just
described. It arises by its superior apex from the
latero-inferior margin of the ultimate tracheal ring,

THE MUSCLES OF THE AIE PASSAGES. 4!)

while its lower apex is inserted into the posterior
extremity of the second half-ring. To do this it must
be evident that it lies obliquely across the larynx.

35. TJie hronchialis anticus has much the same form
as the last muscle, but is fully double its size. It is
crossed at its origin by the broncho-trachealis anticus,
which must be pulled to one side in order to examine it.
We find that its superior apex arises from the last
ring of the trachea, from which point the fibres pass
obliquely forwards to become inserted by a lower apex
into the rim of the arytenoid cartilage of the lower
larynx, and the anterior extremities of the first and
second half-rings.

36, Either aterno-trachealis is represented by a deli-
cate little cord of muscular fibres, which arise on the
side of the trachea beneath the hinder border of
the broncho-trachealis anticus, and pass directly out-
wards, a little downwards and backwards, across
the cavity of the chest, to become inserted on the
inner aspect of the corresponding costal process of the
sternum, rather in advance of its centre. A delicate
baud of fascia spans the insertion of this muscle a few
millimetres in front of its point of attachment. This is
shown in Fig. 19, According to Owen, " this is the
most constant of all the muscles affecting the lower
larynx. It is reckoned by Savart as the sixth })air of
vocal muscles, but not by Cuvier, since it is not directly
attached to any part of the lower larynx, and exists in
birds, as, e.g., the Vulture and Ostrich, in which that
larynx is not developed " {Anat. Verts., vol, ii. p. 224).

The infinite number of changes that the foregoing
muscles can brine about in the form of an organ so con-
stituted as the inferior larynx of the Kaven is, can be
more easily imagined than described. The principal

E

50

THE MYOLOGY OF THE RAVEN,

fact, however, to be kept before us is, that as the form
and tension of its walls vary, the tympanic mcml)rane

Fig. 19. — Author's ski'tcli of Lis dissection of the iiinscnlaturo of the; .lir
passages of a Raven. Shown life-size upon hiteral aspect, cp., de-
tached portion of the costal process of the stevnuni, of the right side,
and turned about.

which stretches across the mesial aspects of the bronchi,
upon the vibrations of which the voice of the bird

THE MUSCLES OF THE AIR PASSAGES. 51

depends, must also be l)rought to different degrees of
tension in its different parts. Striking from the list the
sterno-tracheales, all the muscles we have enumerated
for the true larynx are tensors, as it is evident their
contraction must stretch the tympanic membrane l:)y
lengthening the bronclii to which, as we have just said,
it is attached. On the other hand, if the sterno-
tracheales contract, they must stretch the windpipe, and
thus shorten the bronchi, and relax the tympanic mem-
brane, A glance at Fig. 13, where g is the tympanic
membrane, will at once make these points clear.

Professor Owen found that " in many of the Volitores
there is a single pair of ' broncho-tracheales,' and a
single pair of short ventrieose ' bronchiales.' In Thamno-
pliilus each sterno-trachealis bifurcates to send a small
strip to the lower larynx, and the rest to the side of the
trachea, as nsual. In Furnai^ia the sterno-trachealis is
inserted into the upper end of a long appendage to the
upper bronchial half-ring.

" The Parrot tribe have a single glottis bounded by
a lateral pair of vibratile membranes ; each membrane
connecting together, and occupying the interspace
between, the last tracheal and first bronchial rings.
These have each one margin concave, with the concavity
towards each other, and are movably joined together at
their fore and hind extremities. These half- rings expand,
and stand out from the end of the trachea. A narrow
muscle, 'tensor longus glottidis,' Fig. 105 [14], a, passes
from the side of the trachea to the upper (tracheal)
half-ring ; and, by raising it, makes tense the elliptical
elastic membrane : a broader ' tensor brevis glottidis,' ih.,
b, passes from the lower rings of the trachea to the same
half-ring, diverging to its extremities ; a third narrow
muscle passes from the tracheal to the bronchial half-

E 2

52 THE MYOLOGY OF THE RAVEN.

rings, ih., c, and by approximating them, relaxes the
membrane occupying the elliptical interspace. These
membranes, projecting on each side into or below the
termination of the air-tu])e, leave a narrow chink
between them, through wliicli tlie air passes to and
from the lungs ; and when, in forcible expiration, the
membranes are j)ut into a sufficient state of tension,
they vibrate, and the vocal air is driven along the
trachea through the uj^per larynx, where some modifi-
cation of sound may be made. The tongue of the
Parrot is more fleshy than in most birds. These
structures, concomitant with the single glottis and pair
of vocal folds in the lower or true larynx, relate to the
faculty, so remarkable in these singular birds, of imitat-
ing human speech."

Other interesting variations in these structures are
to be found in certain Cranes, in the peculiar Suij^e
RhynchcBa, in certain Gallinaceous birds, and still more
especially in many of the Anserine fowl.

V. THE MUSCLES OF THE EYE AND EAR.

Undee this head we will notice the folio win 2; : — ■
Those of the eye —

37. Orbicularis palpebrarum. 43. Obliquus inferior.

38. Levator palpebrse superioris. 44. Rectus superior.

39. Depressor palpebral inferioris. 45. Ttectus inferior.

40. Quadratus nictitantis. 46. Rectus externus.

41. Pj'ramidalis nictitantis. 47. Rectus internus.

42. Obliquus superior.

And the single one I propose to describe for the ear —
the

48. Tensor tympani.

To commence the dissection for a study of the muscles
of the eye in the Eaven, we should divide the skin down
to the bone by a circular incision passing completely
around it, a good distance back from the margins of the
lids. Then dissect carefully, reflecting the integument
as we approach these latter, equally all about them.
This is best done by holding its free edge with the thumb
and finger of the left hand, while we dexterously use a
sharp scalpel held in the right, the skin being kept on
the stretch.

37. Tlie orhicidaris i)alpehrarnni is the first muscle we
shall reveal during such a dissection, and its principal

54

THE MYOLOGY OF THE RAVEN.

origin seems to l)e upon the lacrynial and maxillary
bones (Fig. 22, o.p.). Superiorly the circumscribing

.. S

t^r

Fig. 20. — Superior aspect of liead of Raven, with skin on rii^lit side disftocted
away, and the roof of the orhit removed, to sliow muscles of eye witliin.
Life-size, by the author from his own dissections, ob. inf., ohlicpuis
inferior; oh. sicp., obliquus superior; r. int., rectus internus ; r. ex.,
rectus externus ; r. siop., rectus sujjcrior.

fibres become very indistinct, in which locality they are
inserted directly into the ciliary margin. Below, they
are attached to the lower free edge of the tarsal cartilage,

THE MUSCLES OF THE EYE AND EAR. 55

and it is upon this lid that the action of the muscle is
principally exerted, the eye being covered and closed, by
its being drawn upwards over it.

This muscle may now be divided at its principal
origin ; the integument dissected c|uite up to the free
edges of the lids, where we carefully divide it all around,
in order to remove tliese now unnecessary parts. Reflect-
ing the dense tissue above the eye, we behold the
organ in situ.

38. Levator i^alpehrm suj)erioris. — The lower eyelid

Fk;. 2] .—Mesial aspect of eye of Goose, to show muscular insertions (after Owen),
with certain parts removed tliat appear in the original. op. n,, stump
of optic nerve : c, obliquus superior ; d, rectus internus ; /, obliquus
inferior ; b, rectus inferior ; c, rectus e.xternus ; h, pyramidalis ; h\ its
tendon ; q, quadratus ; a, rectus superior.

is the one which principally closes the eye in the Raven,
consequently we find the present muscle but feebly
developed, while, on the other hand, the next to be
described is quite strongly so. The levator palpebrse
superioris arises along a longitudinal line found at about
the middle of the roof of the orbit within. The faiut
and delicate fibres pass directly outwards to find insertion
in the ciliary margin of the upper lid, being best seen
rather on the side towards the outer canthus. As in the

5G

THE MYOLOGY OF THE RAVEN.

Mammalia, the function of this muscle is to raise the
superior eye-lid.

39. TJie depressor palj^ehrce inferioris, as has already
been stated, is easily found. Its fibres arc attached
along the lower margin of the interorbital vacuity.
From this line (Fig. 22) they pass outwards beneath the
eyeball, to become inserted into the lower lid, where they

:^^' ../

:V>V

paZ. tyiiJ^,

T- e*-.

Fig. 22. — Kiglit lateral aspect of skull of Kavcii, to sliow exact origins of the
jirincijial imiscles of the eje, and the tensor tympani. Life-size, by
the author. Lettering as in Fig. 20, with t.t., tensor tympani ; o.ji*. ,
orbicularis palpebrarum; /, lacrynial bone; I. pal. sup. , levator j>al
pebrffi superioris ; d. pal. inf., depressor palpebraj inferioris; r.inj.,
rectus inferior.

antagonize to a certain extent the orbicularis palpe-
brarum, as by their contraction this integumental veil to
the organ of sight, the lower lid, is withdrawn. Owen
tells us that this muscle is also found in the Crocodile,
and I dare say in other Sauropsida. In the Raven it
is the only lid of the three which is reinforced by a

THE MUSCLES OF THE EYE AND EAR. 57

cartilaginous plate, here found between the conjunctival
membrane and the ligamentous layer. It is of interest
to know that in the Strigidce and Caprimulgi the
eyes are closed principally through the depression of
the upper lids.

In common with most other birds, the Eaven possesses
a well- developed memhrana nictitans, which is here
of a pearly w^hite colour, and consequently opaque. As
it closes, it passes obliquely across the eyeball, from its
resting-place in the supero-internal corner, to the one
opposite, a movement effected by two special muscles,
the next to be described (Fig. 21).

40. TJie quadrat Its nictitantis hnds its origin upon the
sclerotic at the upper part of the ball, behind. From this
point the fibres converge as they pass towards the optic
nerve, and are inserted in an aponeurosis which sheathes
the latter above. It is evident that in this course the
fibres of the quadratus pass downwards, inwards, and
backwards, interfering in no way with the action of the
other muscles, have no fixed bony insertion, and do not
stand in the way of the passage of the rays of light.

41. The lyyvamidaUs nictitantis, the remaining muscle
acting upon the membrana nictitans, occurs in the Raven
precisely as we find it in the Goose, and it has been
so well described by Owen for that bird, that I quote his
words. He says the " p^/7'am^G?aZ^s fiictitantis (Fig. 21, h,
of the present work) arises from the low^er and nasal side of
the eyeball : its fibres converge towards the upper part of
the optic nerve, and terminate in a small round tendon
which glides through the pulley at the free margin of
the quadratus ; thus, winding over the nerve, it passes
down to be inserted into the lower part of the margin of
the third eyelid. By the simultaneous action of the
two muscles, that nictitating lid is drawn outward and

58 THE MYOLOGY OF THE HAVEN.

oljliquely downward over tlie fore-part of the eyeball
The tendon of the pyramidalis gains the due direction
for that action Ijy winding round the optic nerve,
and it is restrained from pressing upon the nerve by
the counteracting force of the quadratus, which thus
augments the power of the antagonist muscle, while it
obviates any inconvenience from pressure on the optic
nerve, which its peculiar disposition in relation to that
part would otherwise occasion. The nictitating mem-
brane returns on the relaxation of its muscles, by virtue
of its own elasticity, to the inner corner of the orbit,
where it lies folded when not in use " {Anat. of Verts.,
vol. ii. p. 14;i).

42. Ohliquus superior. — As in the Mammalia, the
movements of the eyeball depend upon the action of two
oblique muscles and four recti muscles (Fig, 22).

The oblique muscles arise close together at the inner
and superior angle of the pars plana, their exact origins
being shown in the figure. Between them passes the
olfactory nerve. The origin of the superior oblique is
situated the more posteriorly, and rather lower, than
that of the obliquus inferior. Its diverging fibres pass
backwards and outwards, forming a pretty, fan-shaped
little muscle, to spread over the upper side of the eye-
ball, their extremities being overlapped by the rectus
superior. In none of these eye-muscles proper do the
terminations of the fibres of insertion reach so far as the
osseous circlet formed of the sclerotal plates ; and it
is evident, further, that, owing to the greatly confined
condition of the eyeball, the necessity for a ligamentous
pulley to assist the action of the present muscle is
obviated.

43. The ohliquus inferior muscle arises, as we have
just stated, close to the last described one (Figs. 20, 22,

THE MUSCLES OF THE EYE AND EAR. 59

oh. inf.). Its fibres pass in exactly the same direction,
differing only in their downward, rather than the upward
inclination, in order to become inserted in the sclerotic
of the inferior aspect of the eyeball, where they are
overlap])ed by the inferior rectus muscle.

The action of these two oblique muscles is too simple
to require a detailed description here. They are the
direct antagonists to each other, and are principally con-
cerned in movements of torsion of the eye.

44. Rectus superior. — The scA^eral origins of the recti
muscles circumscribe the optic foramen, just wdtliin its
free border. The rectus superior and internus surround
its upper half, while the rectus inferior and externus
arise below. The rectus externus arises by two distinct
heads, between which the sixth nerve passes out of
the brain-case through a small circular foramen which
appears to be constant in the Eaven (Fig. 22).

The fibres of the rectus superior diverge as they pass
forwards, upwards, and outwards, the innermost ones
overlapping those of the obliquus superior at right
angles. They are inserted into the sclerotic just within
the margin of the bony circle of the eye (Fig. 20).

45. The rectus inferior muscle is to the rectus superior
exactly what the oblicpius inferior muscle is to the
obliquus superior. Its fibres spreading out like a fan, as
they pass from their origin, are inserted on the lower
aspect of the eyeball, in the same manner as the fibres
of the rectus superior are inserted upon its upper side.

46. The rectus externus seems to be the shortest of all
the recti muscles ; and, as I have already remarked, it
arises by tw^o heads, disposed as shown in Fig. 22. It
also is fan-shaped, and finds insertion, in a w^ay similar
to the two foregoing muscles, on the external, or what
is really the posterior side, of a Eaven's eye.

60

THE MYOLOGY OF THE RAVEN.

47. The rectus internus, after leaving its origin, spreads
out over the mesial aspect of the eyeball, clearing by a
well-devised arrangement the quadratus and pyramidalis,
though in intimate relation with the former : it is finally
inserted, in a manner common with the others of the
group, upon the sclerotic, or the anterior aspect of the

J^rvdUr sid<i^ oj ^^^

Fig. 23.-

-The back of the right eye of a Raven, x 2^, and showing tiie cut
stumps of the recti and ohliqui muscles, whicli, liowever, are drawn
away from the centre to show their proper insertions. The quadratus
nictitantis is represented in a state of semi-contraction, when it lifts the
tendon of the pyramidalis nictitantis off of the optic nerve. Could
the front of this eye now be seen, the mr.mhrana nictitans would be
found about three-quarters drawn obliquely across the eye. The pyra-
viidalis is here contracting then, and it is evident that both its force
and function are augmented by this action of the quadratus.

The curious elliptical osseous plate surrounding the optic nerve,
is also shown in the figure, and I have found it in every Raven's eye
that I have dissected.

eye, or what would be the internal side in the Mammalia.
Were it to act alone, the other muscles remaining passive,
it would so pull the eyeball as to direct the line of vision
to the front.

By way of recapitulation, then, of the nomenclature of

THE MUSCLES OF THE EYE AND EAR. 61

the eye and its appendages in a Raven, we find that its
movements are effected by eleven muscles. The or-
hicularis 2:)alpehrai'um by its contraction closes the
integamental lids. It is antagonized by a feeble levator
"palpehrcB superioris in the upper lid, and a much better
developed depressor jjolpebrcB inferioris in the lower
lid, which latter is the one which really " closes the
eye."

The third eyelid, or memhrana nictitans {nictito, I
wink), is governed in its action by the two muscles, the
quadratus and the pyramidalis, but their operation is
best seen by a study of Fig. 23. AVhen these muscles
cease to act, the nictitating membrane contracts by
virtue of its own elasticity, and again becomes hidden
within the recess of the inner can thus, above the com-
missural point, of the true lids.

The remainino^ six muscles are devoted to the move-
ments of the eyeball itself. Any of the recti, when
acting, antagonize the muscle inserted diametrically
opposite it ; the two obliques similarly oppose each
other, and through the varied traction of the group the
line of vision is directed to meet the will of the bird.
Owing to the close contact of the bony walls of its
chamber, the greater simplicity of the ohliqui, and,
finally, the shortness of all these muscles, the mechanism
involving the movements of the eyeball is not nearly so
complicated a one in the bird, as we find it to be in any
Mammalian form.

Vr. THE MUSCULATURE OF THE EAR.

Upon one of the foregoing pnges I liave already
described the circumconcha muscle, which I believe,
from my dissections, to be a constant one in the Raven.
By its contraction it evidently acts as a"laxator"to
the tympatinm. Careful search in a large number of
ears of this l)ird, however, fails to reveal to me any such
"tensor "as described by Professor Owen. I do find,
though, the following muscle : —

48. Tensor tympani. — If we carefully dissect the
integument about the aural orifice in an old bird of this
species, we find a small fasciculus of muscular fibres that
arise from the inner end of the quadrato-jugal bone, and
the contiguous surface of the quadrate. These pass
beneath the integumental duplicature in the shallow
meatus, to be lost upon the inner surface of the tympa-
num. Now, as the drum is braced in the ear-j)assage
by at least two or three ligamentous bonds, more espe-
cially by a strong one above, it is very evident that, by
the contraction of such a muscle as I here describe, the
ear-drum would be put upon the stretch, and its tense
condition duly effected (Fig. 22, t.t.).

In describing the tensor tynvpani in an Owl (Fig.
\Qi,f.), Professor Owen says : " It arises from a depression

THE MUSCULATURE OF THE EAR. C3

in the basisphenoid, enters the tympanic cavity above
the beginning of tlie Eustachian tube, and, by its inser-
tion into and action upon "the malleus, tends to push the
membrane outward ; it is counteracted by two small
cords extended to the inner walls of the tympanum :
but the muscular character of them is doubtful, and the
ear-drum resumes its normal state when the tensor
ceases to act" {Anat. of Verts., vol. ii. p. 135).

Further investigations in this direction will be very
interesting and quite important. The general text-
books of Claus, Gegenbaur, Jeffrey Bell, Huxley, and
many others at my command, are silent upon this
point. ^

^ Since writing the above I find the following from Professor
Hans Gadow (loc. cit., Bronn's Klassen, vi. Band, pp. 442-445) upon
the musculature of the eyes and the eyelids in Aves : —

" Die Augenmuskeln (Taf. xliii. and iv.).
[Figures not reproduced here.]

" Die 8 Muskelu des Augapfels und des dritten Augenlides ent-
wickeln sich embryologisch aus den Resten der vorderen Kopfsomiten
(vgl. s. 297).

" Der Augapfel selbst wird von sechs Muskeln bewegt, njimlich
von vier ' geraden ' und zwei ' schiefen ' Muskeln.

" 1. M. rectus sujjei'ior s. attollens entspringt fleischig von deni
oberen Rande des Foramen opticum und heftet sich mit breiter, aber
kurzer Aponeurose an den oberen Rand des Uebergangstheiles des
Augapfels. XJeber den Urspringstheil des Muskels iJiuft der R.I.
des N. trigeminus und der N. trochleai'is. Innervation durch den
iV. oculomotorius und zwar durch einen kurzen Ast desselben, der
sogleich noch seinem Eintritt in die Orbita zur unteren Fhiche des
Muskels geht.

" 2. il/. rectus inferior s. deprimens, entspringt fleischig von der
unteren und hinteren Umgebung des Foramen opticum und inserirt
sich mit ziemlich breiter Ausdehnung an dem unteren Rande des
Augapfels. Er wird ebenfalls vom iV. octdomotorius innervirt, und
zwar durch ein Biischel feiner Aeste, die den Hauptstamm nach
Abgabe des Ramus ciliaris verlassen.

64 THE MYOLOGY OF THE RAVEN.

" 3. M. rectus i)dernus s. mediaJis adducens, entspvingt nacli vorn
vom Ansti'itt des Scbnerveu und insei'irt sich am luaenrande des
Angapfels. Er wird medianwarts von der Harder'schen Driise und
vom R. I. JV. tri(/emini, nach oben hiu vom 31. ohliquus superior be-
deckt. Innervirt durch einen Zweig des N. oculomotorius.

"4. M. rectus externus s. lateralis s. abducens, entspringt seit-
lich uud hinten neben dem M. rectus siiperior und inserirt
sich am hinteren Seitenrande des Augapfels. Seitlich von ihm
verlauft der R. II. trigemini. Innervirt durch einen Ast des N.
abducens.

" 5. 31. obliquus sujyerior, entspringtweit nach vorn, von der Eth-
moidalwand ; iiber seinen Urspruug zielit der N. olfactoritis hin,
dicht unter ihn verltivift der R. I. trigemini. Der Muskel liiuft danu
qner iiber den Insertionstheil des 31. rectus internus und heftet sich
sehr breit dicht nach innen, und theilweise unter der Insertion des
M. rectus suj^erior an den Augapfel. Er wird vom A^. trochlearis
innervirt.

" 6. M. obliquus inferior, ein langer dlinner, bandartiger Muskel,
der seitlich vom 31. rectus inferior entspringt und sich medianvviirts
neben demselben inserirt. Nach unten und aussen liiuft iiber seinem
Ursprungstheil der R. superior des N. carotico-cephalicus (s. L. 384)
hinweg. Er wird vom letzten Zweige des jY. oculomotorius in-
nervirt.

" Ausser diesen sechs Muskeln des Augapfels besitzen die Vogel
und meisten Reptilien noch zwei, welche zur Bewegung des dritten
Augenlides dienen.

" 7. 3[. qttadratus menibr. nictitantis, entspringt flach und breit
unter und zwischen den Insertionen des 31. rectus internus, 31.
obliquus superior, und 31. rectus superior. Er ist von trapezoider
Gestalt, indem er an der Basis am breitesten ist und nach dem
Sehnerven hin schmaler wird : letzterer Rand bildet eine aponeur,
otische Schleife fiir die Sehne des 31. pyramidalis. Der 31. quadratus
wild vom N. abduceus innervirt.

" 8. 31. pyramidalis membr. 7iictitantis. Entspringt von der unteren
inneren nasalen Wand des Augapfels, von den Insertionstheilen des
31. obliquus inferior vnid 31. rectus inferior bedeckt. Der Muskel
spitzt sich bald zu, geht median und dorsal vom JV. opticus in eine
runde Sehne iiber, welche dorsalwiirts vom Sehnerven durch die
Schleife des 31. qu,adratus liiuft, dann wieder abwiirts geht und
zwischen den Insertionen des 31. rectus externus und des 31. rectus
inferior auf die Vorderfliiche des Augapfels tritt, wo die Sehne in

THE MUSCULATURE OF THE EAR. 65

die durchsichtige Nickhaut iibergeht. Innervation duich den JV.
abduceus.

'• Die Augenlider [Muskeln].

" Die Bewegung der Augenlider wird durch mehrere Muskeln
bewirkt. Der J/, obicularis, Sphincter, oder Schliessmuskel lauft
kreisformig unter der Aussenhaut der Liderum die Lidspalte herum
und heftet sich an den Lidknorpel an.

" Der M. levator folpebrce entspiingt vom oberen Dache der Augen-
hohle und heftet sich an den ausseren Winkel des oberen Lides.

" Der M. defn-essor palpebrce inferioris ist bedeutend starker als
der vorige, und entspringt in der Tiefe der Augenhcihle. Bei Rhea
bildet der Muskel ein langes, breites Band, welches theilweise vom
Alisphenoid, und lateral ventral neben dem Ursprung des M. rectus
externus entspringt. Es inserirt sich hauptsachlich am hinteren,
unteren Rande des Lidknorpels.

" Bei den meisten Vogeln (Huhner, Schwimm-, Singvogel) ge-
scnieht das Schliessen des Auges durch Heraufziehen des unteren
Lides, und das obere Lid bewegt sich sehr wenig, oder gar nicht,
wie bei den Reptilien. Selten senkt sich auch das obere Lid etwas
durch Contraction des M. orbiculm-is, wahrend das untere sich hebt :
Eulen, Caprimulgus, Tauben. Nur in wenigen Fallen, wie beim
Strauss und bei den Papageien besitzt das obere Lid grcissere Be-
weglichkeit als das untere, wie bei den Sangethieren, Ueberhaupt
ist der Grad der Ausbildung der drei Lidmuskeln ein sehr wech-
selnder.

" Merrem entdeckte beim Adler noch einen ' AugenbrauenmuskelJ
Er ist sehr diinn, entspringt von einer kleinen Hervorragung des
oberen Kandes der Augenhohle und inserirt sich an dem den
Tagraubvogeln eigenthlimlichen Augenbrauenknochen, den er in die
Hohe zu heben scheint.

" Die Innervation der Lidmuskeln geschieht durch den N. oculo-
motorius und durch den K. I. des Trigeininus."

VIL THE MUSCLES OF THE UPPER
EXTREMITY.

In describing tlie muscles of Vertebrates, anatomists
usually, after completing their descriptions of the muscles
of the head, take up next in order the musculature of
the neck and trunk. I have found it to be the case
with birds, however, that by far the most convenient
method of procedure proves to be to take iii^Jirst our
studies and dissections of the muscles of the upper
extremity, and, having completed these, turn our
attention next to the muscles of the lower extremity,
which are to be similarly dealt with, and thoroughly
disposed of, before any of the muscles of the trunk
proper have been examined.

This course gives us the opportunity of clearing away
the very large pectoral muscles of the bird, the heavy
muscles of the thigh, and, finally, removing the limbs
altogether. We then have nothino; left but the much
lighter neck and trunk to deal with, which are easier
handled, and their muscles worked out with far greater
pleasure and satisfaction.

To expose the muscles of the upper extremity we
make a circular incision through the integuments at the
root of the neck. This is met by a linear incision made
on the anterior aspect of the bod}^ following the keel of

THE MUSCLES OF THE UPPER EXTREMITY. 67

the sternum, and following midway between tlie liml).s
of the furcula. Posteriorly, this linear division of the
pectoral integuments is carried well beyond the hinder
sternal margin. Next, make a similar median incision
upon the dorsal aspect of the body, meeting the circular
incision at the root of the neck, and carried posteriorly
as far back as the anterior border of the pelvis. Dissect
away the skin from the left side of the body, and from
the left upper extremity, leaving the right side covered,
in order to keep the muscles moist, and intact for com-
parison, if we find it necessary, with those of the side
we have under examination.

Divide the dermal muscles as we come across them,
during our operation of removing the integuments, at
their middles, allowing the proximal portions to remain,
in order that their relations may be studied.

In removing the integumental fold of the patagium
carefully study the mechanism of the tendons of the
patagii muscles, as well as the muscles themselves. We
also find in this fold other small muscular tendons, which
will hereafter be described. The relations of the pos-
terior patagial fold should also be carefully studied, as
well as the dermal muscles that lie within it. Great
care is necessary when you come to remove the skin
covering the pinion, as it is intimately attached to the
tissues overlying the muscles and tendons beneath it, in
several places, which are liable to be injured by the
scalpel if the removal is too hastily undertaken, or the
skin simply torn off".

THE PECTORAL MUSCLES.

49. The pectoralis major. 51. The pectoralis tertius.

50. The pectoralis secundvis.

This group of muscles is second to none in importance
as compared with any other in the entire muscular

F 2

THE MUSCLES OF THE UPPER EXTREMITY. 69

system of any one of all those birds possessed of the
power of Hight.

49. llie jyecf oralis inajor^ is by far the largest muscle

^ FiirbriDger and Gadow divide the jjectoralis major muscle into
three parts, viz. the pars thoracica, the pars j^ropatagialis, and the
pars ahdomi7ialis .

Omitting the descriptive part, I present the synonymy of these
divisions as recorded by Gadow in Bronn's Klassen ties IVtier-lleichs,
(vi. Band, pp. 241, 242), which is as follows : —

" 73. M. PECTORALIS.

" I. Pars tlioracica.
Depressor alee. Borelli. )

Grand pectoral. Vicq d'Azyr, 1772, p. 623, No. 1.
,, Cuvier.

Gervais et Alix, p. 24; Alix, pp. 399-401.
Pectoralis major. Merrem, p. 152 ; Wiedemann, p. 82.
Grosser Brustmuskel. Tiedemann, § 249.

Meckel, p. 31.5, No. 8; Schiipss, p. 108, No.
15 ; Prechtl, § 35.
„ „ Reid, p. 140 ; Rolleston, p. 4.

De Man, p. 109 ; Watson, p. 80.
Feet oralis major et minor. Selenka, p. 121, No. 46 u. 47.
Pectoralis. Fiirbricger, Morpholorj. Jahrb., v., Monograp)hie ;
Carlsson, p. 26.

" II. Pars p/ropatagialis (Fllrbringer).
Tensor patagii membran. ant. aim. Selenka, No. 48 und 49 (partim).
Sehne vom Pectoralis major zum Tensor brevis patag. ant. De Man,

p. 110.
Verstarkendes Sehnenbiindel vom Pect. major. De Man, p. 111.
Tensor membrance anterioris aloi, Theil a und b. Heusinger, p. 185,

No. 19.
Langer und Kurzer Muskel der vorderen Fliigelfalte. Schopps, No.

2 und 3 (partim).
Spanner das Windfangs. Prechtl, § 69 (partim).

" III. Pars abdominalis (Fiirbringer).
Subcutaneus abdominalis. Wiedemann ; Tiedemann, § 100, No. 5,
,, ,, Watson, p. 55.

„ thoracis. Tiedemann, § 100, No. 4 ; Prechtl, § 71.

70 THE MYOLOGY OF THE RAVEN.

of the system in a Kaven, both as regards weight and
general bulk. It arises from the posterior moiety of the
anterior surface of the sternal body, to the extent shown
in Fio". 25 ; from the entire maro:inal third of the corre-
spending aspect of the keel, as seen in Fig. 24 ; and
finally from the entire outer side of the corresponding
clavicular limb of the furcula, as indicated in Fig. 8.
The fibres from this extensive origin converge, with but
little tendency to overlaj) as we find them in man,
toward the proximal third of the humerus of the same
side, into which they are inserted, by a broad tendon
occupying the entire surface of the palmar aspect of the
pectoral crest. The posterior surface of the pectoralis
major is far more tendinous in structure than its anterior,
and as it passes over the shoulder-joint this tendinous
portion becomes intimately blended with the long head
of the biceps, while near this point it also receives the
insertion of the dermo-humeralis (No. 12), the fan-like
tendon of the latter blending with the broad tendon of
the pectoralis.

In the sternal region the pectoralis major completely
covers the other two pectoral muscles which arise from
the sternal body.

This muscle when contracting depresses the humerus
with great force, and consequently the wing of the bird,
and so forms the chief agent of flight.

In describing the pectoralis major for Ijirds in general,
Sir Eichard Owen says that " This muscle is very long

Ohne Namen erwiilint. Schopss, p. 112.
Dermoliwnieralis. Owen, p. 24.

Fanniculus carnosis (second jmrtion). Eeid, p. 1.39.
Muscle des jmrures. Gervais et Alix ; Alix, p. 401.

,, „ „ Watson, p. 81."

[See the present writei-'s description of the dermo-lnnneraUs in
the present connection.]

THE MUSCLES OF THE UPPER EXTREMITY. 71

and wide in the Natatores generally, but in the Penguin,
its origin is limited to the external margin of the
subjacent pectoral muscle, which is here remarkably
developed. The great pectoral is very long, but not
very thick in the Easores. In the Herons it is shorter,
but much stronger and thicker. Its size is most
remarkable in the Humming-birds, Swallows, and

Fig. 25. — Anterior aspect of Ihe sterr.uni of a Raven, designed to sliow the areas
of origin of the pectoral group of muscles. The subclavius lies beneath
the p. tertvus. Life-size, from the specimen.

diurnal birds of j)rGy, where it is attached to almost the
whole outer surface of the sternum, and its crest,
and has an extended insertion. In the Ostrich its
origin is limited to the anterior and external eighth
part of the sternum, and it is inserted by a feeble
tendon into the commencement of the pectoral
crest of the humerus, to which it gives a strong rotary
motion forwards. In the Apteryx the pectoralis major

V2 THE MYOLOGY OF THE RAVEN.

is represented by two tliin triangular layers of the
muscular fibres attached to the under and lateral part
of the sternum, and converging to be inserted into the
proximal third of the minute humerus" {Anat. of Verts.,
vol. ii., p. 9G).

50. The pectoralis secundiis ' (Figs. 24, 25, 26, 29,
and 30) arises from the remainder of the keel of the
sternum not appropriated by the pectoralis major, as

1 This is the m. supracoracoideus of Professor Hans GadoAv
(Bronn's Klass>tn, vi. Band, p. 246), who has given a very full
description of it, together with the following synonymy, which
I republish below : —

" 74. M. SUPRACORACOIDEUS.

Pectoral moyen. Vicq d'Azyr, 1772, p. 624; Cuvier.

,, ,, Gervais et Alix, p. 24.

Mittlerer Brustmuskel. Merrem, p. 152, No. 2.
Pectoralis minor. Eeid, p. 141.

), r, Nitzsch-Giebel (Upupa, Coracias, Papageien).

Pectoralis minor s. medius. Wiedemann, p. 83.

)> „ ,, ,, Tiedemann, § 250.

,, „ „ ,, Heusinger, § 183.

Zweiter Brustmuskel. Meckel, /System, p. 317, No. 11.
(Pectoralis secundus). Haughton (Emu, Rhea).

,j „ Forbes (Tubinares) ; Weklon (Phoenicopterus

et Leptoptilus).
Deltoides maximus. Schopss, p. 124, No. 21.
Kleiner Brustmuskel (UmroUer des Oberarms). Prechtl, § 40.
Pectoralis medius s. secundus. Owen, Apteryx, p. 289.
Suhclavius. RoUeston, p. 624.

„ Selenka, Bronn, p. 118, No. 44; und Archiv Neer-

land., 1870, p. 48 fp.
De Man, p. 109.
Pectoralis teriius. Jager Literatur, No. 80.
Pectoralis major (partim ?). Eiidinger, p. 89.
Sus-epineux + accessoires du sus epineux. Alix, p. 396-399.
Pectoralis viedius. Watson, p. 82.
Suiyracoracoideus. Fiirbringer, Morphol. Jahrh.^ v. und Mono

graphii ; Carlsson, p. 25."

THE MUSCLES OF THE UPPER EXTREMITY. 73

well as from a contiguous strip of the anterior surface
of the sternal body. It also finds origin from a tendi-
nous expansion stretching between the corresponding
coracoid and the os furcula, and from the anterior
sternal extremity of the lower third of the coracoid
itself (Fig. 24). From these several points of origi]i
the muscular fibres of the second pectoral rapidly
contract as they converge, to pass upward and round
the coracoid to its posterior aspect. As they approach
the shoulder-joint the carneous fil)res become tendinous
to form a rounded and strong cord. This passes through
the canal formed by the scapula, coracoid, and clavicle,
and immediately upon emerging above it is directed
outwards and downwards, to be inserted as a strongs and
somewhat flattened tendon just anterior to the radial
crest of the humerus, on the same side but nearer the
humeral head than the pectoralis major.

It must not be understood from what I have just said
that the fibres of this muscle pass directly to their
humeral tendon, for more strictly speaking they
gradually merge, as they pass in that direction, into a
tendinous sheet which traverses the muscle longitudinally,
and appears as a tendinous line upon its anterior aspect.

Through the leverage gained by the tendon of this
muscle passing through the osseous canal formed by the
bones of the shoulder-girdle, it acts as a levator of the
wing, the humerus being raised by its contraction. And
this mechanism is very prettily effected in spite of the
fact that the chief weight of this muscle is thrown in
favour of depressing the centre of gravity of the bird's
body, a very essential point to be attained, in fact a
positive requirement during flight.

Owen says, " In the Penguins, Guillemots, and Gulls,
this muscle is almost the largest of the three, occuj^ying

74 THE MYOLOGY OF THE HAVEN.

the whole length of the sternum. It is remarkable for
the length and strength of its tendon, which is inserted
so as to draw forwards the humerus with great force.
It is proportionately the smallest in the JRaptoi^es ; and
is very small and slender in the Struthious birds.

"We have already alluded to the use which the
Penguin makes of its diminutive anterior extremities as
water-wings, or fins ; to raise these after making the
down-stroke obviously requires a greater effort in water
than a bird of flight makes in raising its wings in air ;
hence the necessity for a stronger development of the
second pectoral muscle in this and other diving birds,
in all of which the wings are the chief organs of
locomotion, in that action, and consec[uently require as
powerful a developm.ent of the pectoral muscles as the
generality of birds of flight " {loc. cit., p. 97).

Mr. W. A. Forbes says that the pectoralis secundus
"in the Albatrosses is unusually short, and broken up
into four cjuite separate parts, which unite before passing
the shoulder-pulley. In the other Petrels, the muscle
is much more homogeneous, and only separable by
dissection into its various component parts " [Coll.
Memoirs, p. 389).

51. The pectoralis tcrtius^ is the smallest of the

^ Gadow [loc. cit., p. 252) prefaces his description of this muscle
by the following synonymy, he having proposed the name of the w.
coixico-bracidalis posterior for the ])ectoralis tertius : —

" 76. M. CORACO-BRACHIALIS POSTERIOR.

Le petit 2>6Ctoral. Vicq d'Azyr, 1772, p. 625 ; Cuvier.
Kleiner Brustmuskel. Merrem, p. 152, No. 3.
Pectoralis tuinimus. Wiedemann, p. 83.

,, ,, Tiedemann, § 251.

,, ,, Heusinger, p. 183.

Prechtl, § 37.
Coracobrachialis inferior. Meckel, Sysleiii, p, 31i), No. 12.

THE MUSCLES OF THE UPPER EXTREMITY. 75

pectoral group of muscles, and is found to the outer side
of the one last described. As with the first two, its
fibres arise fleshy, and in its case, from an area occupy-
ing the anterior half (in length) of the outer moiety (in
width) of the pectoral aspect of the body of the sternum,
including the broad costal process where it is attached to
the fascia of the suhclavius muscle which lies beneath it.
It also arises from the outer lower third of the
corresponding coracoid process, which latter area lies in
close juxtaposition with the somewhat similar origin of
the second pectoral (Figs. 24 and 25). From these
two origins the fibres of the muscle converge as they
ascend upwards, pass to the outer side of the coracoid
bone, and as they approach the humerus they become

Pectoralis tertius. Schijpss, p. 113, No. 16.

Coraco-brachialis s. Pectoralis tertius s. minor. Rudinger, p. 80.

Coraco-brachialis. Reid, p. 141 ; Gei-vais et Alix, p. 23 ; Alix, p. 393.

Suhclavius. Retzius.

Pectoralis minor s. tertius. Owen, Apteryx, p. 289.

Coraco-brac/iialis longvs. Selenka, p. 114, No. 40 ; De Man,

p. 106 ; Carlsson, p. 26.
Coraco-brachialis internus. Furbringer, Morph. Jahrb. v.
Pectoralis minor. Watson, p. 83.
Coracobracldalis posterior s. internus. Furbringer, Monographic."

Valuable observations are passed upon this muscle by the writer
just quoted in the same connection, and he, after dwelling upon its
anatomy in a number (if bird-groups, declares that, " Er entspricht
nicht dem M. coraco-brachialis des Menschen, der vom Proc. cora-
coideus zum Schafte des Humerus geht. Besser zu vergleichen ist
er einem zweiten 31. coraco-brachialis, der bei vielen Saugethieren
sich am Tuberc. int. humeri besfestigt und bei den Monotremen, die
ein Sauropsiden ahnliches Coracoid besitzen, sehr stark und
vogelahnlich entwickelt ist. Dass er nicht als ein Pectoralis
mi7ior, soudern als Corac. brachial, aufzufassen ist, hat Selenka
(Literatur, No. 121) ausgefuhrt. Bei den Amphibien und Rep-
tilien ist er in dem ebenfalls als Coraco-brachialis beschriebcnen
Muskel zu erkennen" {Joe. cit., p. 2.53).

76 THE MYOLOGY OF THE EAVEN.

tendinous, and finally form a stroncr, somewhat flattened
tendon, which is inserted into the ulnar crest of the
humerus, on the proximal margin of the pneumatic
fossa, at about its middle point.

This muscle assists the pectoralis secundus in elevating
the humerus, when it is brought into action by the
contraction of its fibres.

Professor Owen seems to have found quite a different
origin and insertion in other birds for this muscle, as he
says, " the third pectoral muscle, which is in general
the smallest of the three, arises from the anterior part of
the sternum at the angle between the body and the keel,
and also by a more extended origin, from the posterior
moiety of the inferior surface of the coracoid and the
coraco-clavicular membrane ; it is directed forward,
rising to pass through the scapulo-coracoid trochlea ;
its tendon glides through a sheath, attached to the
capsule of the shoulder-joint, and in some birds to the
OS humero-scapulare ; and is inserted into the radial
tuberosity of the humerus, which it helps to raise. It
is proportionally large in the Penguins and Gulls, but
attains its greatest development in the Gallinaceous
order" {loc. cif., p. 97).

Mr. Forbes, who investigated the myology of the
Tubinares, collected during the voyage of the Chal-
lenger, found the ^^ecfora/^s tcrtius in them to be " in
the form of a broad, thin band, more or less parallel
with the coracoid, occupying the superior half of the
broad space between that bone and the furcula, its fibres
arising chiefly from tlie strong membrane between these
bones, sometimes with additions from the anterior mar-
gin of the coracoid, or from the body of the sternum
close to the middle line " [Coll. Memoirs, p. 390).

Before closing my account of this group of muscles, I

THE MUSCLES OF THE UPPER EXTREMITY

77

must call attention to one other fact in regard to this
third pectoral, and that is this : after having dissected

fa

Fig 26.— Superficial muscles, anterior aspect, of the upper extremity, as far as
the elbow, of a Raven. Drawn life-size by the author from his own
dissections. All of these muscles are in their normal positions, but the
size of the figure did not admit of showing the insertions of the tensor
patagii slips, though their proximal moieties are shown m their proper
positions.

away the pectoralis major, in nearly all specimens, the
outer margins of the second and third pectorals appear

78 THE MYOLOGY OF THE RAVEN.

as though they were continuous, or a prolongation of
the same oblique line ; this is indicated by the shading
across the belly of the pectoralis tertius in Fig. 27. In
very muscular subjects, however, this muscle does not
terminate at this line, but very delicate fibres pass
beneath the fascia so as to cover an area of oriixin as
indicated for the pectoralis tertius in Figs. 24 and 25,
where its full extent or limit is shown for a very power-
fully-developed, old muscular male bird.^

DORSAL MUSCLES OF THE UPPER EXTREMITY.

Several of the muscles to be described under this head
are generally spoken of by anthropotomists as " nmscles

1 In certain birds there is a small muscle in the axillary region,
which I have failed to discover in a Raven, and its place seems to
be in some ways replaced by the dermo-ulnaris muscle. Mr. Garrod
describes it in the following words for Cltauna ; he says the
^^ Expansor secundariorum is the name which it is my habit to
employ for a very small and peculiar triangular muscle arising from
the quills of the last few (generally two or three) secondary
remiges at the elbow. Its remarkably long and slender tendon,
which frequently traverses a fibrous pulley on the axillary mai-gin
of the teres muscle, runs up the arm side by side with the axillary
vessels and nerves to be inserted in the thorax, into the middle
of a tendon which runs from the inner side of the middle of
the scapular element of the scapulo-coracoid articulation to near
the middle of the thoracic border of the sterno-coracoid articulation,
at right angles to it when the fore-limb is extended. This arrange-
ment being found very well diiferentiated in the Storks may, for
the sake of convenience, be termed Ciconine " (see Collected
Scientific Memoirs, 1881, p. 32.3, and plate showing this muscle
in Chauna, No 16, where it is marked e.s. ; or the same plate in the
P.Z 8. of 1876, No. xiv.). The author has as yet never examined
any of our American Herodiones for this muscle.

Since writing this footnote, nearly three years ago, my oppor-

THE MUSCLES OF THE UPPER EXTREMITY. 79

of the back," but, as I have already decided above, it is
my intention to consider all those muscles which find

tunities to examine into the literature of this subject have much
improved, and (in Bronn's Klassen des Thier-Reichs, vi. Band,
p. 258) I tind a very excellent description of this muscle by
Professor Gadow, too long, I regret to say, to reproduce here. This
will not apply, however, to the synonymy which he there presents,
and which reads as follows : —

" 78. M. METAPATAGIALIS.

Una 2^0Ttion du grand dorsal. Vicq d'Azyr, 1772, p. 632, No. 5.
Tensor memhravue jjosterioris alee. Wiedemann, p. 85 ; Tiedemann,

§267.
,, ,, ,, ., Biidinger, p. 91.

Selenka, No. 50.
Spanner der hinteren Flughaut. Meckel.
M. plicce alaris posterioris. Schopss, p. 79, No. 1.
M. coracobrachialis hrevis (pt.). Milne-Edwards, Ossem. fossil.
M. expaiisor stcundariorum. Garrod, F.Z.S., 1876, pp. 193, 194,

und 199.
„ ,, „ Forbes (Tubinares, p. 29).

M. 7)ietap)atagialis. Fiirbringer."

—(June 14, 1889, R.W.S.)

In October 1887, I published in The Journal of Comparative
Medicine and Surgery (New York) an essay in which was reviewed
the muscles used in the classification of Birds (see No. 124 of Bibl.
at end of the present volume), and there I made the following
comments, and said that " the expansor secundariorum (Fig. 35 his,
Exp. Sec.), although of insignificant size, is a muscle that has proved
of no little value as a classificatory one. Garrod spoke of it as the
Ciconine chai-acter, as it was so well developed in the Storks. It
occurs in quite a large number of groups of birds, as the Gallince;
the Ducks, Geese, and Swans j the Rails, Plovers, and many others.
While ' in the majority of the Gallinaceous birds the expansor
secundariorum, with the normal origin from the secondary quills,
has a different method of insertion, which has led Mods. A. Milne .
Edwards to describe the muscle in the Common Fowl as a part of
the coraco-brachialis (brevis) in his superb work on fossil birds '
(Garrod).

"Professor Sutton alludes to this muscle in the following interest-

80 THE MYOLOGY OF THE RAVEN.

origin or insertion upon any of the bones of the arm, or
tlie shoulder-girdle, as muscles of the upper extremity.
For we cannot clear the subject of this part of its
anatomy until these are disposed of, and described.

52. The latissimus dorsi. 60. The subcLivius. '

53. The trapezius. 61. The coraco brachialis.

54. The rhomboideus. 62. The teres minor.

55. The coraco-humeralis. 63. The levator scapulte.

56. The scapulo-humeralis. 64. The thoraco-scapularis.

57. The supraspinatus. 65. Tlie subscapuhiris.

58. The teres et infraspinatus. 66. The serratus parvus anticns.

59. The serratus magnus anticus.

ing way. He says, ' Every student of human anatomy must have
experienced a certain amount of curiosity when he dissected for the
first time the plantaris muscle ; this strange structure sinks into in-
significance when compared with the celebrated ambiens of the bird's
leg, or the tendon of the femoro-caudal in the Lacertilia. Of all
strange muscles, the one known as the expansor secundariorum
(Garrod) in the bird's wing, stands pre-eminent. It is a small
triangular muscle, arising from the quills of the last few secondary
remiges at the elbow. Its remarkably long and slender tendon,
which fiequently traverses a fibrous pulley on the axillary margin
of the teres muscle, runs up the arm side by side with the axillary
vessels and nerves, to be inserted in the thorax into the middle of a
tendon, which runs from the inner side of the middle of the scapular
element of the scapulo-cox'acoid articulation, to near the thoracic
border of the sterno-coracoid articulation, at right angles to it when
the fore-limb is extended.

" ' In the ducks and geese, among the Anseres, the tendons under
consideration, when they enter the thorax, run towards one another
and join (after having expanded out), in the middle line in front of
the oesophagus, and behind the trachea.

" ' My investigations into the morphology of this tendon induce me
to believe that it is the representative in the bird's wing of the
coraco-brachialis longus of mammals, and the long brachial ligament
of man ' {Ligaments, their Nature and Morphology, p. 33).

" This will prove a vei-y interesting muscle indeed to search for in
the various forms of bird life in our own United States avifauna."

THE MUSCLES OF THE UPPER EXTREMITY. 81

52. The latissimus dorsi,^ next to the dermal muscles
of the region, is the most superficial muscle of the dorsum.

1 From Gadow {loo. cit., pp. 226, 227) I obtain the following
synonymy of the latissimus dorsi : —

" 68. M. LATISSIMUS DORSI.

Grand dorsal. Vicq d'Azyi-, 1772, p. 631.

,, „ Cuviei" ; Geivais at Alix, p. 21.

Hinterer anziehender Armmuskel und Ruckwartszieher des Arms,

Merrem, p. 153, Nos. 7, 8.
Latissimus dorsi und spinalis brachii. Wiedemann, pp. 84, 85.
Latissimus dorsi. Tiedemann, § 252.
,, ,, Heusinger, p. 183.

„ ,, Meckel, System, p. 313.

„ „ Schijpss, p. 103.

„ „ Reid, p. 141.

,, ,, Nitzsch-Giebel.

,, ,, Owen, Ajjteryx, p. 288.

„ „ Selenka, p. 120, No. 45.

De Man, p. 109.
,, ,, 'Pvivhrmgev, Morphol.Jahrh., V.

, ,, Haswell, Proceed. Lin. Sac. New Sotith Wales,

1880, p. 306 ff. ; 1883, p. 115; id., Journal
Anat.Phys., 1883, p. 219.
,, ,, Gadow (Pterocles, Tauben), Fi-oc. Zool. Soc,

1882, p. 321.
„ ,, "Watson, p. 87.

„ ,, Weldon, p. 641 ; Carlsson, p. 19.

Breiter Riickenmuskel (Rlickwiirtszieher des Oberaims). Prechtl,

§42.
Latissimus dorsi + teres major. Haughton {Dromoius, p. 496).

Rhea, p. 503 e, Fig. 35.
Pars metapatagialis (Fiirbringer) = pt. Tensor membrance j^ostei'ioris
alee und M. plicce alaris jwsterioris der Autoren."

This author splits the latissimus dorsi up into " I. Der vordere
Theil ; II. Der hintere Theil ; III. Die Pars metapatagialis.'^
These several divisions of the muscle as thus divided receive
quite careful description at his hands, while for the physiology of
.the muscle he republishes the observations of Prechtl, as he does
those of Fiirbringer for the comparative morphology.

G

82 THE MYOLOGY OF THE RAVEN.

It is well developed in a Eaven, and as in the majority
of birds, divided into two distinct slips. As a whole,
however, it is a thin, fan-shaped muscle, or more
properly speaking, a triangular one, with its apex at
its insertion upon the humerus, and its hase at its
origin, attached to the vertelu'al spines.

The anterior slip arises from the outer edges of the
superior margins of the neural spines of that vertebra
that bears the last pair of free rd^s, and the one next
succeeding it, whose ril)s articulate with costal ribs ;
the posterior slip arises from the similar margins of
all the following neural spines of the verte1)r8e of the
dorsum, which are four in number. This latter origin
partakes very largely of a thin fascia-like character.
From this combined origin, the fibres of the intimately
connected slips rapidly converge as they pass directly
to the humerus; they enter between the deltoid, the
scapular head of the triceps, and the remaining heads
of the latter muscle, to be inserted upon the shaft of
tliat Itone, on its anconal aspect, just within the
maro-iu of the radial crest, as a flat, ril)bon-like
muscle in this division of its course (Fig. 27).

53. The traj^ezius ^ lies immediately beneath the

^ Attention is invited in this connection to Gegenbaur's Anat. des
Mensehen, 1883, p. 311. The following synonymy I take from
Gadow (Bronn's Klassen, vi. Band, p. 217), who describes the
trapezius under the name of the m. rhomboideus superjickdis : —

" 65a. M. KIIOMBOIDEUS SUPERFICIALIS.

Trajyezoide. Vicq d'Azyr, 1772, p. 630, No. 1.
Trapeze. Cuvier ; Gervais et Alix, p. 21.
Aufzieher des Schulterblatts. Merrem, p. 154, No. 9.
Kappenmuskel s. Cucidlaris. Wiedemann, p. 84.

„ ., ,, Tiedemann, § 242.

„ ,, „ Schopss, p. 90.

Selenka, p. 107, No. 32.

Tig. 2 7

~<'-^

^^^l^jKon c^.j,iy ttZrvajrls.

Fig. 27.-

FiG. 28.
Fig. 29.-
Fig. 30,
Fig. 31.

Anconal aspect of right humerus of <i Raven, designed to show the areas of
oiigin of the muscles attached to it. The os Immcro- scapular e is drawn in the
position it normally occupies during life. The deltoid muscle is attached to it.
-A proximal view of the head of the same humerus.
■The same ; palmar aspect.
The same ; radial aspect.

-The same ; ulnar aspect. Figures are all life-size, and drawn by the author
from his own dissections.

G 2

84 THE MYOLOGY OF THE RAVEX.

latissimus dorsi for its posterior two-thirds, while
the anterior or remaining third is superficial. The
muscle is quite well-developed, consisting of a flat
layer of fil)res, of an oblong figure, which arise just
l)elow the latissimus dorsi, on the neural spines of the
two vertebrae that bear free ril_)S, and the next two .
succeeding; vertebras of the dorsum. From this orio;in
the fil)res pass directly across to the scapula to become
inserted upon its mesial border to the extent shown
in Figs. 8 and 24. The jjosterior moiety of the trape-
zius overlies the anterior portion of the rhomljoideus
muscle, but the direction of their fibres is difterent,
as those of the latter pass backward and outwards,
while, as I have said, those of the present muscle pass
directly i^Cross to their insertion (Fig. 33).

54. The r/umihoideus^ in man and most mammals

Kappeiimuskel s. Cucidlaris. De Man, p. 103.

Oberriachlicher Eiu- unci Ruckwiirtszielier. Meckel, Sijstem, p. .30G,

No. 1.
Trapezms. Watson, p. 76 ; Weldon, p. 641.
lUiOiuholdes 2it. [cuctdUiris). Flirbringer, Morjjh. Jahrh., v.
lihoiiihuides sujierjiclalis. Flirbringer, Morph. Jalirb., xi., und

Mo7iograp]de.
Rhomhoideus inferior. Weldon, p. 641."

Gadow further adds (Bronn's Klassen, p. 218) that " Dieser
Muskel, der bei den Vcigeln passend als M. spini-scapularis bezeich-
net werden kann, ist als ein oberflaclilicher Theil des Rhom-
boideus der Siiuger zu betrachten, daher nicht, yfie bisher fast
allgemein geschehen, mit dem Cucullaris zu verwechseln. Zuriick-
f uhriiug auf Anipliibien und Reptilien ist schwierig, da er wohl mit
dem wahreii Cucullaris eine noch nicht differenzirte Masse bildet,
jedoch liisst er sich nach Flirbringer mit der gleichnamigen Bildung
der Crocodile homologisiren."

^ " 656. M. RHOMBOIDEUS PROFUNDUS.

Ii/iomhoide. Yicq d'Azyr, 1772, p. 630; Cuvier ; Gervais et
Alix, p. 21.

THE MUSCLES OF THE UPPER EXTREMITY. 85

is represented by two distinct muscles, wliile in birds
it is invarialjly a single one. It is absent altogether
in the Apteryx. Here in the Eaven, it arises from
the neural spines of the first four dorsals, or at least
from the spines of the leading four vertebrae, that
possess completed ribs. This origin lies l)eneath that
of the latissimus dorsi and the trapezius muscles,
where they overlap it. The fibres pass backwards
and outwards to insert themselves upon the posterior
third of the mesial border of the scapula, commencing
anteriorly about where the trapezius leaves oft', the
overlapping of these two muscles, in all the specimens
I have examined, being at their origins only.

On the upper side of the shoulder-joint we find in
the Raven two very slender muscular slips, passing
from the shoulder-girdle to the humerus. They both
act as levators of the brachium, and perhaps too, to
a certain extent, counteract each other in pulling the
humerus forwards and backwards.

Rautenmuskel (rhomboid eus). Wiedemann, p. 82.

,, ,, Meckel, System, p. 307.

„ „ Schopss, p. 93.

Selenka, p. 108, No. 33.

„ „ De Man, p. 104.

„ „ Watson, p. 77.

„ „ Carlsson, p. 17.

Rliomboideus major et minor. Tiedemann, §§ 243, 244.
Rhomboides pt. (rhomhoideus). Fiirbringer, Morph. Jahrh., v.
Rhomhoideus profundus. Fiirbringer, Monoqraphie.
Rliomboideus superior. Weldon, p. 641." (See Gadow in Bronn's
Klassen des Thier-Reichs, vi. Band, pp. 218, 219.)

"Der Rh. profundus stellt eine secundare Differenzirung des
Serratus profundus dar, die bei manchen Vogeln (z. B. mehreren
Ratiten) noch keine voile Selbststandigkeit gewonuen, bei anderen
(z. B. Pici und Capitoniden) nocli weiter geliende Sonderungen und
Zerfallbildungen erkennen lasst " (Fiirbringer).

86 THE MYOLOGY OF THE RAVEN.

As a rule, these muscles are not fully described in
works devoted to the myology of birds, and until such
time as the homologies in myology are better known,
I have Ijestowed the following names upon these two
little muscular slips : —

55. TliG coraco-htmieralis} the smaller and more
anterior of the two, arises from the outer side of the
head of the coracoid, above and rather posterior to the
origin of the long head of the biceps (Fig. 24, cJi.).
Its filu'es form a delicate, subcylindrical cord that passes
to the head of the humerus to become inserted upon
its palmar aspect, beyond and to the inner side of the
insertion of the j)ectoralis major (Fig. 29).

5G. The scapnlo-humeralis '^ (Fig- 8) arises from the

1 Flirbi'inger defines this muscle as the coracohrachialis anterior
s. externus, in his magnificeut work upon the morphology of the
class Aves ; and Gadow presents us with the following synonymy : —

" 75. MUSCULUS CORACO-BRACHIALIS ANTERIOR.

Le muscle qui correspond au coraco-bracJiial. Vicq d'Azyi-, 1773,

p. 568, No. 5.
Deltoideus inferior. Schopss, p. 122, No. 20 (partim).
Deltoideus minor. Heusinger, p. 183, No. 6.
C oraco-hrachialis jyroprius s. pectoralis niedius. Riidinger, p. 89.
Deltoideus minor. Selenka, p. 117, No. 43.
Coracohracldalis anterior s. externus. Flirbringer (Joe. cit., p. 251)."

2 According to Gadow (loc. cit., p. 234) this muscle represents
the deltoideus minor of Fiirbringer, and he gives the following
synonymy of it : —

" 70. M. DELTOIDEUS MINOR.

Le petit relevetir de Vhumerus. Vicq d'Azyr, 1773, p. 567.
Levator humeri. Tiedemann, § 255 (?).

,, ,, Heusinger, p. 183, No. 7.

Deltoideus externus. Schopss, p. 120, No. 19.
Kleiner deltaformiger Muskel (Vorwiirtszieher des Oberarmes).

Prechtl, § 39.
Deltoideus minor. De Man, p. 108.

THE MUSCLES OF THE UPPER EXTREMITY. 87

inner side of the neck of the scapiihx just within the
origin of the deltoid ; its fibres form a narrow, flat-
tened ribbon, that passes over the top of the shonkler-
joint, parallel to the upper margin of the larger portion
of the deltoid, to become inserted on the palmar aspect
of the humeral head, directly between the insertions
of the pectoralis major and secnndus (Fig, 33, s.h.).

Professor Owen describes for the Apteryx a suh-
scapularis muscle as arising " from the anterior part
of the inner surface of the scapula, and is inserted
into the ulnar humeral tuberosity." I might have
considered this the same as the muscle described
above as my scapulo-lnmieralis, but the eminent
anatomist just quoted says further that the suh-
scapularis is divided into two portions by the pec-
toralis minor.

Unfortunately, I have not Professor Owen's drawings
of the myology of Apteryx before me at the present
writing, but it is difficult for me to understand from
his description in The Anatomy of Vertebrates, how
the muscle he calls the suhscapularis can be " divided
into two portions by the pectoralis ininor,^' when he
says of the latter that " A muscle, which may be
regarded either as a portion of the pectoralis mino-r,
or as the analogue of the suhclavius muscle, arises
from the anterior angle of the sternum, and is in-
serted into the external margin of the sternal extremity

Deltoidevs minor. Fiirbringer ; Carlsson, p. 20.
Accessoire coracoidien du sus-spineux. Alix, p. 399 (?)."

Considerable confusion has attended the identification of this
muscle, and the deltoideus viinor of Selenka (Bronn's Klassen,
vi. Band, p. 117) is not taken by Gadovv to be the muscle he
describes under that name, and the pi-esent writer is satisfied that
the d. viinor of Gadovv is the muscle described above as the
scajndo liumeralis.

88 THE MYOLOGY OF THE KAVEN.

of the coracoid bone " {Anat. Verts., vol. ii. p. 95).
Of course it is hard for me to judge, as I say, in
the absence of Professor Owen's figure, but if the
fibres of this suhclavius yruscIq have the same direction
as his pectoralis minor, then it is hard to see how
they coukl divide the suhscajyularis in two portions,
which pass between the scapula and the humerus.
(For the way these muscles occur in the Eaven, see
No. 60 et seq.)

57. The supraspinatus ^ (Figs, 24, 27, and 34)
is a thin, flat muscle of a triangular form, its base
being represented by its origin, and its apex by its
insertion. It arises from the superior surface and outer

1 " Recht stark entwickelt ist der Muskel bei Corvus," says
Gadow, in speaking of the sup-aspiaafus, which he describes in
Broun's ThierReichs as the scapuli-humeralis anterior, with the
subjoined synonymy (vi. Band, ji. 235) : —

"71a. M. SCAPULI-HUMERALIS ANTERIOR.

L' humero-scajiulaire. Vicq d'Azyr, 1773, p. 569.
Schulterarmmuskel. Wiedemann, p. 86.

Prechtl, § 41.
Humerosca'pularis parvus. Tiedemann, § 257.
,, ,, ,, Heusinger, § 184.

Su}iraspinatus,o([ev Teres minor ? Meckel, System, p. 312, No. 6.

„ „ „ „ Schopss, p. 107.

Teres minor. Reid, p. 142 ; Nitzsch-Giebel.
Subscapidaris {inferior posterior). Macalister, p. 16.
Infraspinatus. Retzius, Selenka, p. 113, No. 38.
De Man, p. 106 ; Carlsson, p. 19.
Teres minor s. Infrascapularis. Gurlt, p. 2 1 .
Le petit rond. Gervais et Alix, p. 22.
Supraspinatus. Riidinger, p. 86.
,, Watson, p. 85.

Weldon, p. 643.
Infraspinatus. Fiirbiinger, Morph. Jahrh., v.
Scapido-humeralis anterior. Fiirbringer, Monographie."

THE MUSCLES OF THE UPPER EXTREMITY.

89

i-i. ^OThgiog

Fig. 32. — Anterior aspect of a number of muscles of the upper extremity, and
more particularly designed to show in situ the 2}<^c^oralis secundus
and tcrtius, and the biceps. The pcctoralis major has been almost en-
tirely removed, a, the end of cut tendon oi pcctoralis secundus; b,
its humeral stump ; c, the humeral stump of the pecto7'alis major.
Life-size, from an adult female Raven, by the author, from his own
dissections.

rounded margin of the scapula, posterior to the origin
of the long head of the triceps, for about the anterior

90 THE MYOLOGY OF THE RAVEN.

tliird of the hlade of tlie bone. Its filn-es converg-e to
pass directly to tlie corresponding liumerus, to become
inserted upon tlie anconal aspect of the bone, just within
the border of the lower part of the pneumatic fossa.,
below the origin of the deltoid, and l)etween the forks
of the external and internal heads of the triceps, wdiich
really seems to divide to give it space to make fast
to, as shown in the figures.

58. The teres et infraspinatus^ is the name I have

^ As will be seen iu my Preface to the present volume, I was
far removed from nearly all liooks, and from all libraries and
anatomists, when it was written, and when this muscle was
primarily described by me. So when the synonymy of these parts
as collected by Gadow recently came to my hands, it was with no
little interest that I noted that the muscle now under consideration,
as proves to be the case in quite a number of others I was obliged
under those circumstances to bestow names upon, had already
received the name I coined for it by Tiedemann, by Heusinger,
and by Schopss.^(I4th June, 1889.)

The following is its synonymy by Gadow (Bronn's Klassen des
Thier-Reichs, vi. Band, pp. 236, 237) : —

" 715. M. SCAPULI-HUMERALIS POSTERIOR.

Sus-scajndaire. Vicq d'Azyr, 1772, p. 631. »

Schulterblattmuskel. Merrem, p. 154 ; Prechtl, § 38.
Ober-Schulterblattmuskel. Wiedemann, p. 87.
/Su2)ra-sca2)ula7-is {Infraspinatus + teres 7najor). Tiedemann, § 256.

„ „ ,, „ „ Heusinger, p. 184.

Sujyrascajjularis. Gurlfc, p. 21.

Untorgriitenmuskel. Meckel, System, p. 312, No. 5.
Trifrasjnnatus s. teres major. Schiipss, p. 105.
Irtfras2jinatus. Reid, p. 141 ; Owen, Apieryx, p. 288 ; Watson,

p. 86.
Teres major. Retzius ; Selenka, p. 113, No. 37 ; De Man, p. 105.

,, ,, Furbringer, Morjih. Jahrh., v. ; Carlsson, p. 1*J.

Teres Diinor. Macalistei', p. IG.
Le grand rami. Gervais et Alix, p. 22.

„ ,. „ Alix, p. 394.
Scajmlolmmeralis jwsterior. Fllrbringer."

THE MUSCLES OF THE UPPEH EXTREMITY. 91

bestowed upon tlie present muscle, as it seems to com-
bine tlie tei-es major and the sivpraspinatus of such
vertebrates wherein these muscles are found se])aTate.
Even in old muscular subjects among the Ravens, one
is almost led to l)elieve that the two can be fairly dis-
tinguished, the division being indicated by a somewhat
broader dividino- fascia.

The teres et injfrasjmiatus possesses fully four or
five times the l^ulk of the swpraspinatus, and is the
largest muscle, by all odds, that is attached to the
shoulder-blade (see Figs. 24, 27, and 34).

It arises from the superior surface and the outer
margin of the hinder two- thirds of the scapu]a, extending
in fact between the posterior termination of the supra-
spinatus to the apex of the bone. Its fibres rapidly
converge as they pass forwards and outwards to their
point of insertion upon the corresponding humerus.
This is found upon the anconal surface of that bone
at about the middle of the free ulnar margin of the
pneumatic fossa ; the external head of the triceps
forking to allow this muscle room to attach itself
(Fig. 27).

Owen says in the Apteryx that it " is inserted into
the ulnar tuljerosity of the humerus, wdiere it is closely
attached to the capsule of the shoulder-joint."

The major portion of both of these last-described
muscles He beneath the latissimus dorsi, the supra-
spinatus being the less covered of the two (Fig. 33).

Chief among the uses of these muscles is the protection
they afi'ord the shoulder-joint, and acting as braces
they greatly assist keeping the humerus in its shallow
socket. After the wing has been extended, they, by
their contraction, assist to close it again by drawing
the humerus towards the side of the Ijody. They arc

92 THE MYOLOGY OF THE RAVEN.

also useful in some of the movements of tlie bracliium
during flight ; and it must be evident to meet any
of these ends, the supraspinatus is the one possessing
the least power, while the te7^es et wfraspinatiis, from
its greater size and more favourable origin, is the more
powerfully endowed.

59. TJie serratus ma gnus anticus ^ is composed of

^ In dealing with the serratl group of muscles in birds, Gadow
divides his m. serraUis superficialis s. t/ioraci-scapularis into three
parts, viz. : — [. the pars anterior, II. the pars posterior, and III.
the y^ars melapjatagialis. His work (Bronn's Klassen, vi. Band,
pp. 221-224) presents descriptions of these several divisions, to
Avhich the student is referred, and records the subjoined synonymy,
herewith republished : —

"666. M. SERRATUS SUPERFICIALIS S. THORACI-SCAPULARIS.

*' I. Pars anterior.

Costo-scapndaire. Vicq d'Azyr, 1772, p. 629 ; Cuvier.

Costo-scajjularis inferior. Wiedemann, p. 87.

Costo-scajmlaris. Tiedemann, § 247.

Kleiner Brustmuskel oder vorderer sageformiger Muskel. Meckel,

Systeni, p. 308, No. 7 ; Schijpss, p. 96.
Serratu8 magnus anticus {first portion). Owen, Ajjteryx, p. 288.
Serratus anticus {j't.). Selenka, p. 110, No. 35.

,, ,, ,, De Man, p. 105; Carlsson, p. 18.

Grand dentele anterieur. Gervais et Alix, p. 20.
Serratus anticus minor. Watson, p. 78.
Serratus suj^erficialis, pars anterior. Fiirbringer.

'* II. Pars posterior.

Sous-scapulaire. "Vicq d'Azyr, 1772, p. 632.

Grand dentele. Cuvier.

Eiickwartszieher des Schulterblattes. Merrem, p. 154.

Serratus. Wiedemann, p. 87.

Serratus magnus. Tiedemann, § 246.

„ ,, Fiirbringer, Morpih. Jahrh., v.

„ „ Weldon, p. 641.

Grosser, vorderer Sagemuskel. Meckel, System, p. 308, No. 6.
„ „ „ Schopss, p. 94.

THE MUSCLES OF THE UPPER EXTEEMITY. 93

three strong digitations, each of considerable bulk.
The first of these slips arises from the outer aspect
of the second true dorsal rib (counting from before
l^ackwards) rather above the base of its epipleural
appendage. The other two slips arise from similar
points on the next two succeeding ribs, just above
the origin of the derrtio-ulnaris. The extremities of
these digitations of the serratus magnus anticus, are
strongly inclined to partake of a tendinous character,
and the first two may be more or less attached to the
interpleurapophysial membrane. They converge as

Serratus viagnus anticus {iniddle and jwsterior portion). Owen,

Apteryx, p. 288.
/Serratus anticus {x>t.). Selenka, p. 110, No. 35.

„ „ . De Man, p. 105.

Grand dentele jmsterieur. Gex'vais et Alix, p. 20.
Serratus anticus major. Watson, p. 77.
Serratus superficialis, j>clts j^osterior. FUi-bi'inger.

" III. Pars metapatagicdis. Fiirbringer.
(pt.) Tensor menihranoi p^ostcrioris ahv.. Wiedemann; Tiedemann ,

§ 267.
„ „ „ „ Riidinger, p. 91.

„ „ „ . ,, Selenka, p. 123, No. 50 ;

Carlsson, p. 18.
(jit^ M. plicm alar is posterior is. Schiipss, p. 79."

If the reader will refer to my dermo-ulnaris (No. 11 of the
present work) he will find that that muscle represents the pars
metapatagialis of the above synonymy ; so that the synonyms here
will probably stand thus —

SnUFELDT. GaDOW.

No. 59, Serratus magnus anticus — I. Pars anterior.

No. 64, Thoraco-scapuloi (in pt.)")

1^0. Q&, Serratus parvus anticus >= II. Pars p)Osterior.
(in part) )

No. 11, Dermo-ulnaris = \\1. Pars metapatagialis.

04 . . THE MYOLOGY OF THE RAVEN.

they 2>a,s.s iipwni'ds and slightly forwaixls, and are
inserted on the inferior surface of the apex of the
corresponding scapula (see Figs. 24, 34, and others).
60. Tlie suhclavius ^ muscle arises from the entire
outer surface of the costal process of the sternum and
the contiguous outer surfaces of three or four of the
hsemapophyses, in which situation it is largely over-
lapped by the pectoralis tertius. Its outermost sheath
of fascia is attached to the inferior external maroin
of the corresponding coracoid Ijone, l)ut its short and
oblique muscular fibres pass over this to be inserted
and fill the fossa that is found at the lower third of
the posterior aspect of the coracoid. When this muscle
contracts it simply pulls the coracoid very slightly
outwards, the bone sliding along upon the articulation
of its sternal bed. The more important function of

^ Both Fiirbringer and Gadow describe this muscle as the steryio-
coracoideus, while originally it was considered by Tiedemaun and
Schopss to be the suhclavius.

Gadow presents us with the following synonymy for it (Bronn's
Klassen des Thier-Reiclis, vi. Band, pp. 224, 225) : —

" 67. M. STERNO-CORACOIDEUS.

Le davicidaire court. Yicq d'Azyr.
Riickwiirtszieher der Schllisselbeine. Merrem.
M. clavicularis e:rternus. Wiedemann.
M. suhclavius. Tiedemann, § 248.

,, ,, Schupss.

Ohne Namen, oder vielleicht kleimer vorderer Siigemuskel. Meckel,

System, p. 308, No. 8.
Pectoralis minor. Ketzius.

Suhclavius s. pectoralis minor. Biidinger, p. 89.
Serraius anticus minor. Owen, Apteryx, p. 288.
Coraco-sternalis. Selenka, p. 11], No. 36.

„ ,, De Man, p. 105 ; Carlsson.

Sterno-coracoidetis. Furbrioger, Morph. Jahrh., v., xi., und Mono-

(jraphie.''

THE MUSCLES OF THE UPPER EXTREMITY. 95

the muscle consists in keeping the coracoid firmly in
place and preventing its dislocation during violent action
of surrounding muscles (see Figs, 8 and 25).

61. The coraco-hrachialis ^ (Figs. 8 and 27) is a
long fusiform muscle that arises by a delicate tendon

^ My dissections of the coraco-hrachialis in Corvus convinced me
that it had the same origin and insertion described for it by Owen
[Anat. Verts., vol. ii. p. 97) ; but I find that Gadovv, who calls the
muscle the subcoracoideus, finds a different arrangement of things,
for he says that, " Bei Corvus, Garrulus, und Paradisea entspringt
ein Theil von Binnenseite und dem Vorderrander des Sternum und
von der Tnnenflache des benachbarten Coracoidtheiles ; ein anderer
Theil kommt vom Proc. fui-cularis des Coracoids und der Scapula.
Beider Theile Sehnen inseriren sich vereinige dicht am Caput
humeri etwas proximal von der Sehne des Subscapularis " (Bronn's
Klassen des TJiier-Eeivhs, vi. Band, p. 239). Gadow agrees with
Owen in that the coraco-hrachialis is missing in Struthio and small
in the Apteri/x.

He presents us with the following synonymy for it (loc. cit.,
p. 238) :-

" 72rt. M. SUBCORACOIDEUS.

Souclavier iiiterne. Vicq d'Azyr, p. 628.
Vorderer anziehender Armmuskel. Merrem, p. 153.
Ohne Namen. Meckel, System, p. 320, No. 13.
Coraco-hrachialis siqjerior. Schopss, p. 115, No. 17.
Coraco-hrachialis brevis. Milne-Edwards ; Selenka, p. 115, No. 41.
„ „ ,, De Man, p. 106 (2. Portion) j Carlsson,

p. 18.
Coraco-hrachialis. Owen, A-pteryx, p. 289 ; Watson, p. 84 ; Gurlt.
L'accessoire coracoidien du sous-scapulaire. Gervais et Alix, p. 23.
Coraco-hrachialis externus. FUrbringer, Morph. Jahrh., v.
Suh-coraco-scapularis (Pars coracoidea) — M. suhcoracoideus. FUr-
bringer, Morph. Jahrh., xi., und Monographic.'^

Authors appear to describe two portions of this muscle, of which
my coraco-hrachialis is one ; Selenka says of his Coracohrachialis
hrevis (No. 41) that " Die alteren Beschreibungen dieses Muskels
sind siimmtlich mangelhaft und verwirrt, viele sonst ausfiihrliche
Arbeiten nennen ihn nicht eiumal. Cuvier spricht nur allgemein

96 THE MYOLOGY OF THE RAVEN.

from a small circular origin .situated at the postero-
mesial aspect of the shaft of the coracoid, immediately
above the fossa which harbours the insertion of the
subclavius. Its fibres pass almost directly upwards
parallel with the coracoidal shaft, and as they near
the head of the humerus they again become tendin-
ous, and are finally inserted on the top of the ulnar
tuberosity, on a diminutive subcircular space, common
to it and the insertion of the teres minor.

Owen tells us that " This muscle is wanting in the
StrutliionidcB, is of small size in the Heron and Goose,
is much more developed in the Raptores and many
Natatores, especially the Penguins, and attains its
greatest relative size in the Rusores, where it arises
from almost the whole of the coracoideum."

62. Tlie teres minor '^ arises fleshy from the under
side of the anterior tip of the scapula, the clavicular
process of the bone, to the extent shown in Fig. 8.
The fibres, forming a chunky, little muscle, pass
directly outwards, behind the coracoidal head, and
beneath the neck of the scapula. Clearing these two
bones the fibres rapidly converge to form a small
tendon, which is intimately fused with the tendon of the
coraco-brachialis, and consequently makes the same

von zwei Muskeln, die am Os coracoideum entspringen und am
Humeruskopfe sich inseriren ; das siiid die beiden hier als
C oracohrachiales beschriebenen Muskeln. Tiedemann fiihrt einen
Deltoideus minor und Levator humeri auf, dei zum Theil wenigstens
dem Coracobrachialis brevis entsprechen. Auch Gurlt, Wiedemann,
Merrem, Aldrovandi erwahnen den Muskel nicht " (Bronn's
Klussen des TJiier-Reichs, vi. Band, p. 115).

^ Possibly this muscle corresponds to the subscapularis intermis
of Gadow's " 72b. M. subscapularis," and attention is invited to
the muscle I desci-ibe below as the sid>scapularis (No. 65), and the
footnote under it.

THE MUSCLES OF THE UPPER EXTREMITY.

97

insertion on the top of the nhiar tuberosity of the
corresponding liumerus.

To a limited degree, the coraco-brachiaJis and teres

Fig. 33. — Superficial muscles of the upper extremity in the region of the shoulder
and dorsum of a Raven, s, scapula ; a, anterior slip of the latissimus
dorsi ; p, its posterior slip. The head of the coracoid can be seen where
the mesial extremities of the patagii muscles pass over it ; s.h., the
scapulo humeralis muscle. The extensor metacarpi radialis longus
muscle is indicated by the letters e.m.r.l. Life-size, by the author-,
from his own dissections.

minor assist the pectorals in their action in the down-
ward stroke of the humerus, and so the wing, but they
also assist in keeping that bone in its shallow glenoid
cavity.

H

98 THE MYOLOGY OF THE RAVEN.

Ill all of the works at my disposal I fail to find
a description of tliis teres minor muscle, and consequently
have taken the responsibility of naming it myself.
In its origin, insertion, and action, it seems to corre-
spond quite closely with the teres minor of many of
the higher vertebrates, in arising near the neck of the
scapula and being inserted upon the tuberosity of the
humerus.

Mivart says that the teres minor may coalesce with
the infraspinatus, as in the Two-toed Ant-eater. The
minor may lie larger than the major, as in the Horse
{Ehm. Anat., p. 326).

63. The levator scapidcB^ arises hj two strong,
fleshy digitations, one from each of the lateral processes

1 This is the serratus jn'o/undus of Fiirbringer and Gadow, and
the latter presents us with a very full description of it {loc. cit.,
pp. 220, 221), together with the following synonymy : —

" 66«. M. SERKATUS PROFUNDUS.

Anzieher des Schulterblattes. Merrem, p. 1 54.
Costo-scapularis superior. "Wiedemann, p. 87.
Levator scapulce. Tiedemann, § 245.

Selenka, p. 109, No. 34.
,, „ De Man, p. 104 ; Fiirbringer, Morpli. Jahrh., v. ;

W^atson, p. 79 ; Weldon, p. 641 ; Carlsson,
p. 17.
Schulterheber. Meckel, System, p. 307.

„ Schopss, p. 97.

L'angulaire. Gervais et Alix, p. 20.
Serratus profundus. Fiirbringer.

" Riidinger bemerkt ganz richtig, dass in Folge seines Ursprungs
von den Rippen dieser Muskel eine wesentliche Abweichung vom
Levator scapulae der Saliger bietet, da er bei letzteren von den
hinteren Zacken der QuerfortsJitze der ersten 4 Halswirbel ent-
springt. Er entspringt dagegen ziemlich genau dem M. collo-
scapularis der Reptilien, wenn man die Lange des Vogelhalses in
Betracht zieht. Der Serrat. prof. (Levator scapulae der Autoren)

THE MUSCLES OF THE UPPER EXTREMITY. 99

of those vertebrae that support the free pairs of ribs ;
the fibres slightly converge as they pass directly back-
wards to become inserted into the middle third of the
inferior aspect of the blade of the corresponding scapula.
When this pair of muscles contracts they will evidently
pull the scapulae directly forwards, and with it, of
course, the entire shoulder-girdle, which will articulate
like a hinge-joint at the coraco-sternal junction.

Owen says that " the levator scapula3 arises hj digi-
tations from the pleurapophyses of the last cervical,
and the first two dorsal vertebrae ; it is inserted into
the posterior part of the dorsal edge of the scapula,
which it pulls forwards. In the Apteryx it seems to
be the most anterior portion of the series of fasciculi
composing the serratus magnus anticus." It will be
seen from this description that both its origin and inser-
tion is very diff"erent in the Kiwi-kiwi from what it is
in the Eaven.

64, The tlioraco-sccqiidains ^ is the name I here be-
stow upon that muscle in Corvus cor ax which arises
from the outer aspect of the lower half of the ultimate
free rib ; from the middle of the outer side of the next
succeeding rib, or true dorsal, and its epipleural

zeigt in Gegensatze zu den Reptilien namentlich bei den Carinaten
eine gewisse Vereinfachung, die z. Th. dadurch entstanden ist, dass
ein Theil von ihm sich als besonderer Muskel (Rhomboideus pro-
fundus) differenzirt undabgetrent hat " (Gadow quoting Fiirbringer,
p. 221).

1 Probably the muscle here described is the ^jars jjosterior (in
part) of Gadow's ni. serratus suiyerficialis s. thoraci-scapularis, the
synonymy of which is given under the serratus maynus anticus of
the present memoir, No. 59 (which see).

The remaining part of the pars j^osterior of Gadow's w. ser7-atus
svperjicialis, appears to be herein represented by my serratus joarvus
anticus (No. 66), and the reader's attention is also invited to that
muscle.

H 2

100 THE MYOLOGY OF THE KAVEK

appendage ; and finally by another digitation, the
largest of the three, from a like origin on the onter
snrface of the following rilj. These three digitations
form, a broad, flat muscle lying close against the parietes
of the thorax. Its fibres converge and pass upwards and
forwards, but as they enter the scapulo-thoracic spaci;
they become converted into a broad, thin, flat tendon,
which, passing between the two divisions of the sub-
scapularis, is inserted on a line, covering the junction
of the middle and anterior thirds of the outer margin
of the corresponding scapula (Fig. 24). This muscle,
when the ribs are fixed, draws down the scapula to the
side of the chest, but when the shoulder-girdle is
fixed by the opposing set of muscles, it draws up the
ribs to which it is attached, and thus increases the
capacity of the chest, and performs an important
function in the act of inspiration.

From what I quoted from Sir Richard Owen, above,
it would seem that this muscle corresponded wdth his
'pectoralLS minor, and if we be permitted to compare
the two, it seems to correspond even with the pectoralis
minor in a man, but a far greater amount of study will
have to be undertaken than has been done up to the
present time, before we can say much about such
homologies.

Mivart says of the pecto]'aUs minor that " the
smaller pectoral is a much less constant muscle than
the large one, being very frequently absent. Even in
animals closely allied to man {e.g. many Apes) it is in-
serted into the capsular ligament of the humerus instead
of into the coracoid process. It may form one with the
pectoralis major, as in birds" {Elem. Anat., p. 326).
It will be seen that this last statement of this authority
cannot be reconciled with what we have found to l^e the

THE MUSCLES OF THE UPPER EXTREMITY. 10 1

case in tlie Raven and many otlier birds. Any way we
may regard it, however, the fact still remains that in
many vertebrates, we have a muscle that passes from
the middle outer surfaces of some of the leading ribs,
to become attached to some part of the shoulder-girdle, or
scapular apparatus, and that this muscle is an important
one in inspiration. We may call it the pec^^07'(x/^s
minor, or the thoraco-scapularis, as I have in the present
instance, as we may please.

65. Tlie suhsccqmkiris ^ is one of the largest and most
powerful of the muscles in the Corviche, and is well

^ See No. 62 of the present memoir in this connection (the teres
minor), and the muscle here described appears to be the I. {suhscap.
externus) of Fiirbringer and Gadow, or in other words, as near as
I can make it out, my teres minor + (my) suhscapularis = the
suhscapularis (with internal and external part) of Fiirbringer.
Gadow (Bronn's Klassen, vi. Band, pp. 240, 241) presents us with
the following valuable study of the subject : —

" 72J. M. SUBSCAPULARIS.

Sousclavier externe. Vicq d'Azyr, 1772, p. 628.
Vorderer anziehender Armmuskel. Mensem.
Sous-scapulaire. Cuvier ; Gervais et Alix, p. 22.
Unter-Schulterblattmuskel {suhscapularis). Wiedemann, p. 89 ;

Schopss, p. 128, No. 22 ; Meckel, System, p. 321.
? Deltoideus externus. Schopss.
/Suhscapularis. Reid, p. 142.

,, Riidinger, p. 87.

,, Macalister (superior posterior), p. 1 6.

„ Haughton, pp. 497a, 504.

Gurlt, p. 22.
„ Selenka, p. 113, No. 39.

„ Watson, p. 85; Carlsson, p. 18.

C oraco-hrachialis hrevis (I. Portion). De Man, p. 106.
Suh-coraco-scaj^ularis [Pars scap)ularis iiiterna + externa) — Suh-
scapularis internus et externus. Fiirbringer.

" Der Suhscapularis ist in seinem Ursprungstheil vcm unteren
Rande des Scapulo-humeralis posterior durch die Insertion der Pars

102 THE MYOLOGY OF THE RAVEN.

developed in the present subject. It arises from tlie
anterior and outer half of the bone, including the
margin l)etween the superior and inferior surfaces. Its
fibres rapidly converge as they pass towards the
head of the corresponding humerus, and just before
reaching that bone they become converted into a small,
though strong, subcylindrical tendon, which is inserted
on top of the ulnar tul:)erosity close to the combined
tendons of the coraco-hracJiialis, and the teres minor.
The flat tendon of the thoraco-scapularis divides the

anterior M. serrati superf. getrennt. Dieser Theil uud der Scapul.
humeral, post, liegen dorsal, der Subcoracoideus ventral vein
Sabscapularis.

"Der ganze Muskel besteht nach Ptlidinger's deutlicher Beschrei-
bung aus zwei durch die Pars anterior M. serrati superficialis
getrennten Abtheilungen, von denen die obere [Sabscap. extern.)
vom lateralen Eande des Schnlterblattes, die untere {Subscap. intern.)
von der ventralen Flache der Scapula entspi'ingt. Vereinigt ziehen
beideum den hinteren unteren Theil des Schultergelenkes herum
und heften sich mit einer kurzen Sehne an den unteren hinteren
Theil des Tuberculum mediale s. minus humeri, nahe an dem
Ansatzpunkte der Gelenkkapsel. Wahrend dieser Muskel aus der
Fossa axillaris heraustritt, nimmt er ein accessorisches kleines
Biindel vom Os coracoideum auf.

" Die speciellen Verhiiltnisse sind folgende.

"I. Der obere Theil [Suhscapul. externus) ist gewuhnlieh der
kleinere und entspringfc bei Haliaetos, Colwmba, Fsittacus, Corvvs,
Garrulus vom zweiten Fiinftel, bei B^iteo vom erstea Drittel, bei
Falco tinnunculus, bei Ardea, Anser und bei den meisten Wasser-
vogeln von der basalen Halfte des lateralen oder vorderen Randes
der Scapula, greift auch auf deren Aussen-, oder auf die Innenflache
liber. Bei Fidica atra ist der Ursprung auf das erste Yiertel,
bei Sphenisciden und Hiihnern auf das erste Fiinftel oder noch mehr
beschrankt.

" II. Der untere, grossere Theil [Suhscapid. intevnus) reicht vom
oboren Theile des Coracoids an gewohnlich auf die erste Halfte
{Haliaetos, Corvus, Columba, LejjtojJtUus) , oder auf die ersten zwei
Drittel (Buteo, Falco)."

Fig. 34. — Oblique left lateral view of the second layer of muscles of the upper
extremity, being those of the dorsum and arm, of a Raven. The
heads of the ribs have been disarticulated from the vertebrae, and the
latter drawn up with a dissecting hook and chain, the better to show
the trcqiezitts and rhomhoidcufi muscles. The os humero-scapu/are has
been entirely freed of both the deltoid and ligamentous attachment,
though left in exactly its normal position in this bird. Life-size, by the
author, from his own dissections.

104 THE MYOLOGY OF THE RAVEN.

posterior moiety of the belly of tliis muscle into two
portions, covered by separate sheaths of fascia, but in
the smaller and anterior division the fil;)res again come
together and are included in the same sheath.

In the Apteryx this muscle has very much the same
origin and insertion that it has in the Eaven. Ac-
cording to Mivart it " is generally in Mammals much
as it is in the human sul)ject. It may be considerably
smaller, however (as in Cetaceans) ; and by a singuiar
exception it may (as in the Echidna) exclusively arise
from the external surface of the scapula " [Eleu.
Anat., p. 326).

The subscapularis is a powerful rotary muscle of tLe
head of the humerus, and also an important auxiliaiy
in retaining that bone in its shallow facet, and 'prcr
venting displacement.

66. The serratus 2:)arvus anticus ^ is a thin, flat musch
covering the outer parietes of the chest. It arises by
three rather extensive digitations from the outer surfaces
of the first free rib, and the next two succeeding ones
that connect with the sternum, above the origins of the
seri^atus tnagnus anticus and the thoraco-scapularis.
The fi.bres in a flat sheet of fascia pass upwards and back-
wards, to become inserted in the inferior margin of the
corresponding scapula, along a line to the outer side of
the insertion of the rhomboideus, as indicated in Fig. 8.
In the Penguins this is the largest of the muscles of the
scapula.

In the Eaven at least, the levator sccqnilce, the
serratus parvus anticus, and the serratus magnus
anticus, all really belong to the same system or series,

^ See footnotes and synonymy under muscles described in the
present work as the serratus magnus anticics and the thoraco-
scajmlaris (ISTos. 59 and 64).

THE MUSCLES OF THE UPPER EXTREMITY. 105

which amono; most Mammals is known as the serratus
7nagmis muscle. In certain Reptiles, as the Chameleon,
it may be in several detached parts, whereas in Meno-
poma it is of a very diminutive size.

THE TEKSOR PATAGII MUSCLES.^
67. The tensor patagii loogus. 68. The tensor patagii brevis.

Under this head I will separately describe two extra-
ordinary little muscles peculiar to the musculature of the
pectoral extremity of the vast majority of birds.

^ Referring to 'Bronn' s Klassen des llder-Reichs (vi. Band, pp. 253,
254) we find a very clear and succinct account of these muscles by
Gadow, who records the following synonymy for them ; they being
by him both described under

"77. M. PROPATAGIALTS.

Le grand {et le jjetit) extenseur de la membrane externe de Vaile. Vicq

d'Azyr, 17 73, p. 568, No. 3 und 4.
Langarmiger Muskel. Merrem, p. 156.
Tensor membrance anterioris alee. Wiedemann, p. 85 ; Tiedemann,

§ 267; Heusinger, p. 185, No. 19.
Spanner der vorderen Flughaut. Meckel, pp. 337-345.
Spanner des Windfangs. Prechtl, § 69.

Langer Muskel der vorderen Flligelfalte. Schupss, p. 82, No. 2.
Kurzer ,, ,, ,, ,, ,, p. 86, No. 3.

ilf. accessorius ad bicipitem. Nitzsch.
Tensor longus + et brevis (patag. membran. anter. aim). Selenka,

No. 48 und 49.
Tensor patagii longus + brevis. De Man, p. 110, No. 15 und 16;

Garrod, Liter., No. 56 ; Froc. Zool. Soc. 1876, pp. 508-512, und

Taf. 48-51 ; Forbes, Tubinares, pp. 25-28, und Taf. iv., mit

ausgezeichneten Abbildungen ; Carlsson, pp. 20 und 21.
Tensor p)0'tagii longus. Watson, p. 88.
Tenseur marginal de la membrane anterieure de Vaile. Gervais et

Alix, p. 23.
Cleido-metacarpien — Tenseur marginal. Alix, p. 402, No. 1.
Cleido-epicondylien — Tenseur moyen. Alix, p. 403, No. 2.
M. proimtagialis. Fiirbringer."

lOG THE MYOLOGY OF THE RAVEN.

Tliey have been noticed by every anatomist wlio has
at any time interested himself with the morphology of the
group,while the lamented Garrod successfully pointed out
the fact as to how they might be used, and really were
valuable factors in taxonomy. In the Collected Scientific
Papers of this last authority we find them alluded to in
the following words, viz.: — "In the triangular patagium
of the wing of the bird the tendons of two muscles are
to 1)6 found. One is that of the tensor j^cttagii longus,
which forms the supporting cord of the free margin of
the meml)rane itself. The second is that of the tenso)-
2?atagii hrevis, which courses j)arallel wdth the humerus,
not distant from that bone, to the muscles and fascise
of the forearm." Professor Garrod follows these remarks
hj a full and clear description of the peculiarities of
these muscles in the vast numljer (200+) of birds which
he had especially dissected and examined in relation
to this particular structure.

His highly valuable contril)utions are rather too long
to incorporate in the present connection, Init they will
l)e found in the Proceedings of the Zoologiccd Society
of London for 1877 (pp. 506-19) and in that rare
and imperishable work, his Collected Scientific Papers,
edited Ijy the late Mr. W. A. Forljes, another labourer
in the same field, whose loss to us it is hard to over-
estimate.

Professor Owen's account of these fleshy slips runs
thus : —

" A remarkable muscle, partly analogous in its
origin to the clavicular portion of the deltoid, but dif-
ferently inserted, is the extensor pliccB alaris, and
forms one of the most powerful flexors of the cubit.
It is divided into two portions, of which the anterior
and shorter arises from the internal tuberosity of the

THE MUSCLES OF THE UPPER EXTREMITY.

107

liumeriis ; the posterior and longer from the chivicular
extremity of the coracoid bone. In the Ostrich and Rhea,
however, both portions arise from the coracoid. The

SuhcjLa^y-us Xm'

S vi-6 sC'Cif' t-i-ict^' ViS

Fig. 35. — Left oblique view of the body of a Raven, dissected to sliow the deep
muscles about the shoulder-joint. The coraco-brachialis has been
pulled out by a hook and chain, and the scapula has been rotated out-
wards to sliow the muscles beneath. Other parts have also been placed
in a favourable position to expose the muscles to be shown ; somewhat
diagrammatic. Reduced one-third, from dissections by the author.

posterior muscle sends down a long and thin tendon
which runs parallel with the humerus, and is inserted,
generally by a bifurcate extremity, into both radius and

108 THE MYOLOGY OF THE RAVEN.

ulna. The anterior muscle terminates in a small tendon
which runs along the edge of the aponeurotic expansion
of the wing. In this situation it becomes elastic ; it
then resumes its ordinary tendinous structure, passes over
the end of the radius, and is inserted into the short
confluent metacarpal. It combines with the jDreceding
muscle in bending the forearm ; and further, in conse-
rpience of the elasticity of its tendon, puckers up the
soft part of the fold of the wing " [Anat. Verts., vol. ii.
p. 98).

And Mivart says, when speaking of the trapezius,
that in Bats a long slender segment of this muscle
may pass along the upper margin of the wing mem-
brane from the occiput to the distal jihalanx of the
pollex.

" In the Flying Squirrel {Pteryomys) a similar mus-
cular band goes to the rudimentary pollex, luit it
springs from the zygoma, and is therefore rather a
modification of the platysma myoides than of the
trapezius ; as it is also in Galeopithecus.

" In birds an analogous and similarly slender muscle
goes to the pollex or to a sesamoid at its base, but this
muscle is often an ofl"shoot from the pectoralis major,
though it may contain fil^res from the deltoid or from
the biceps — showing in what diverse ways a similar
want may be supplied" [EJem. Anat., p. 320).

I present the results of the observations of these
several eminent authorities, more particularly to show
how very different are the relations, origins, and in-
sertions of these two little muscles in the class birds,
and how well accurately recorded dissections of them
would repay the labours of the investigator. Several
years ago I showed how the distal extremity of the
tensor patagii longus was attached to the os j^i^omiyiens

THE MUSCLES OF THE UPPER EXTREMITY. 109

ill many of our American Hawks and Owls (Bull.
Nutt. Oniitho. Club, vol. vi. No. 4, October, 1881). I
also showed how among these latter birds, when the
wing w^as extended the tendon of the tensor patagii
longus raised the os proniinens, and thereby increased
the extent of the alar superficies {loc. cit., p. 200).
In foregoing paragraphs of the present work, I have
already described the dermo- tensor patagii muscle,
and how it acts as an auxiliary to the tensor patagii
longus.

Before entering upon our detailed descriptions of the
tensor jpoAcigii muscles, I desire to say a word here as to
how they axe best studied in general. This I do from
the fiict that in the future there will no doubt be many
dissections made of these muscles, as they have been
proven to be of no little classificatory value in birds, as
I have already remarked.

First, it may be as well to add, that in certain birds
another small muscle occurs in this region ; it has been de-
scribed as the bicipital slip to tliepcdagium (Figs. 35 his
and 35 ter, B. slip). This is a fleshy fasciculus of muscle
that is difiJ'erentiated off from the anterior surface of
the biceps, and passing between the cutaneous folds
of the patagium becomes inserted into the tendon of
the tensor patagii longus at about the middle of its
course.

Garrod states that " the presence or absence of this
muscular fasciculus is a very constant character among
closely allied birds." He not only found it in tlie Capri-
mulgi, but also in Plovers, Cranes, Gulls, Auks, and
some few other groups. It is absent in the Corvidce. In
Fig. 35 ter, I present its appearance as it occurs in our
Mourning Dove, a bird I especially dissected to show it
as an illustration in the present connection.

110

THE MYOLOGY OF THE RAVEN.

Professor T. Jeffcry Parker describes this muscle for
the Common Pigeon {Zootomy, j)- 251) as the tensor
patagii acccssorius, and snys, " Its anterior Ijorder is

^^. 35 tcr.

Fig. 35 5?'s.— Axillary muscles of the side of Gallimi.la chJoropus. By the author,
after Garrod. £xp. Sec, cxpaiisor secundariorum ; Pcct. 1., great
pectoral muscle ; JJ. slip, hiceps slip ; Bi, biceps ; T, triceps ; and
other letters as in former figures.

Fig. 35 tcr. — Outer asjiect of the axillary muscles of the right arm in a specimen
of tlie Mourning Dove {Zenaidura macroura). From the specimen, hy
the author, and after his own dissections. Letters as before.

connected by fascia with the tendon of the tensor longiis,
and from its posterior border a long stout tendon is
given off which passes outwards, soon becoming parallel

THE MUSCLES OF THE ITPPER EXTREMITY. Ill

to the tendon of the tensor longus, and having a com-
mon insertion with it."

If this last tendon be present in our wild pigeons, it
is very feebly developed and consequently easily over-
looked, I did not detect it in the Dove above alluded
to, and must believe it was absent in that particular
specimen. Wherever I examined it, it has invariably
agreed with Garrod's description of it.

Now what I have said above in reference to the dissec-
tion of these muscles in the Eaven, applies wdth equal
truth to any bird. Let us choose for our purpose an
oriole, for instance, and in that species the tensor patagii
longus, tensor patagii brevis and dermo-tensor patagii
all occur. Having secured a good specimen of some such
bird, pluck it perfectly clean, being careful, in removing
the feathers of the wings, that you do not tear the skin
(Fig. 35 quat.). Now with a small, sharp dissecting
scalpel make an incision just through the integuments
and no more, along the line which I have indicated by
the letters inc. in Fig. 35 quat., and then carefully and
completely reflect this skin-Hap in both directions until
all the superficial muscles of the arm and forearm are
exposed, even to the tendon which extends from the
shoulder to the wrist [tp. I.), in the free margin of the
duplicature of the skin in which these muscles are found.

By practice we soon learn the best way of doing this,
and in the case of all ordinary sized birds, our left hand
soon learns how to hold the wing in such a way that the
parts to be examined are kept on the stretch, thus faci-
litating our examinations and study, which latter may
now be undertaken.

Professor Garrod chose the wing of Icterus vulgaris
to illustrate the arrangement of the patngial muscles in the
Passer es, and it will be seen by an examination of his

112

THE MYOLOGY OF THE RAVEN.

figure given below (Fig. 35 quin.) that lie found in that
species the tendon of the dermo-tensor 2)(:(^taf/ii merging

Fig. 35 quat. — The plucked body of a medium sized passerine bird, with its
pterylosis not shown, but with the patagial muscles of tlie arm in
blai:k, and supposed to be seen through the skin ; dt. p., dermo-teusor
pata^ii ; ip. 1-, tensor patagii longus ; and tp. h., the tensor patagii
brevis ; inc., the line of incision to be made in order to expose them for
examination. Drawn by the author.

the

with the tendon of the tensor patagii longus, near
shoulder-joint, much in the same way as the present

THE MUSCLES OF THE UPPER EXTREMITY.

113

writer found it to exist in the Eaven and other Corvidce.
In this connection the reader should refer back to
the account of the denno-tensoi' ^ja^a^/Z^-' in the present
work (No. 6).

We wouki naturally expect this to be the case, as
Icterus and the Eaven are members of nearly related
families.

Fig. 35 quin.—yiew , from the outer side, of the muscles of the patagiiim of
the left wirig of a passerine bird. Troiipial {Jderus vulgaris). (After
Garrod.)

Among some other passerine birds, however, the
tendons of the tw^o muscles just mentioned do not
blend with each other in the manner we have described,
but remain quite distinct as far as the carpus. This
is the case in our Purple Martin (Progne subis), as I
have shown in Fig. 35 sex.

114

THE MYOLOGY OF THE RAVEN.

Some of the most com^^licated and interesting con-
ditions assumed by these muscles are to be seen among
such groups of sea-fowl as the Tuhincires, and Forbes
presented us with drawings and descriptions of many
of these in his contributions to the Proceedings of the
Zoological Society of London.

Now in the Raven we find a still different con-

FiG 35 sex. — Same view and corresponding parts of the same wing of a passerine
bird, Purple Martin, Prognc subis (by the present writer), t.p. I., tensor
patagii longiis ; tp. b., tensor patagii brevia ; dt. p., derrao-tensor
patagii ; c.in.i\l., extensor metacarpi radialis longus; t., triceps; b.,
biceps ; -S'. ^. , secondary remiges. (Nearly x 2.)

dition of affairs from some of those alluded to above,
in the arrangement of these two little muscles and
their tendons, so far as their origins and insertions
are concerned, which I will proceed now to describe.
Ql. The tensor 2^o.tagii longus (Figs, 8, 26, 32, and

THE MUSCLES OF THE UPPER EXTREMITY. 115

33) muscle arises by a tendon common to it and the
tensor j^citagii hrevis from the siipero-mesial line of
the head of the corresponding clavicle. This tendon
is flat and thin, being closely approximated to the
head of the coracoid and the outer angle of the
'pectoralis major muscle. The tensor 2)atagii longus
soon becomes muscular, however, after leaving its
origin, to form a slender, spindle-shaped belly a1)out 3 '5
centimetres long. From its distal apex then a tendon
is given off, which runs in the marginal duplicature of
the patagium, and with which filiro- elastic membrane
it is intimately connected. This tendon increases in
calibre, and for more than its entire middle third it
becomes in turn of a fibro- elastic structure. Finally
passing over the end of the radius and to the inner
side of the tendon of the extensor metacarpi radialis
lo7}gus, it is attached to the os radiale of the carpus,
and the fascia which l)inds down the other tendons on
the anterior aspect of the wrist-joint.

68. The tensor ijatagii hrevis arises from the head
of the clavicle, as described for the last muscle, by a
common tendon. The muscular portion of this short
tensor of the patagium is very considerably broader and
longer than that of the long tensor of the membrane
in question. It also overlies the former to a certain
extent near their common origin, and is pointed
downward towards the elbow instead of outwards
towards the carpus. Its tendon is straight, slender, and
nearly of uniform calibre. At about 1'5 centimetres
from the distal end of the humerus it meets the
extensor metacarjn radialis longus muscle of the
antibrachium. There its tendon bifurcates, and the
shorter and distal-projecting bifurcation is soon merged
into the fascia of the muscle just alluded to, of the

I 2

116 THE MYOLOGY" OF THE RAVEN".

forearm ; while its loiio;er and more tendoii-like fork
runs back with the tendon of the extensor metacarpi
radialis longus, to Ijecome inserted just heloiv the
tubercle of the external condyle of the humerus, the
insertion of the extensor itself monopolizing the outer
aspect of the tubercle in question (Fig, 33).

The action of these muscles is well described in the
quotations of the several eminent authorities I have
incorporated with my own descriptions above,

MUSCLES OF THE BRACHIUM.

We come now to take up those muscles of the
ujDper extremity which more properly belong to the
humeral reoion, and are in most instances inserted
into the bones of the forearm. Under this division of
the muscles of the upper extremity, I will notice the
following : — ■

69. The biceps, 7L The triceps.

70. The deltoid. 72. The brachialis anticus.

G9. The biceps ^ is a large and powerfully developed
muscle in the Raven. To study it satisfactorily we

^ For fur-ther and important studies of the biceps muscle in birds,
see Gadow [loc. cit., p. 261), who records the subjoined synonymy : — -

"79. M. BICEPS BRACHri.

Le biceps. Vicq d'Azyr, 1773, p. 570; Cuvier.
Zusammenleger des Fliigels. Merrem, p. 155, No. 3.
Zweikopfiger Armmuskel. Wiedemann ; Prechtl, § 46.
Biceps. Tiedemann, § 258 ; Heusinger, p. 184.

„ Meckel, System, p. 322 ; d' Alton, p. 22.

Riidinger, p. 99 ; Selenka, p. 124, No. 51 ; De Man, p. 111.

,, FUrbringer ; Carlsson, p. 26.

Biceps brachial. Gervais et Alix ; Watson, p. 102 ; Alix, p. 405.
Flexor avtibracliii longus s. biceps brachii. Schopss, p. 135, No. 24."

Gadow divides the muscle into two parts — I. m. bice2)s brachii,
and II. pars pjropatagialis (compare footnotes under the account of
the tensor patagial muscles of the present work).

THE MUSCLES OF THE UPPER EXTREMITY. 117

must carefully remove the common integuments of
the arm and proximal moiety of the forearm ; divide
the tendons of the patagii muscles ; and dissect away
all the superfluous adipose tissue, fascia and vessels.
The biceps will then be seen to be a large sub-
fusiform muscle, occupying the anterior aspect of the
arm. Its proximal third is composed of a broad,
flat tendon covering the top of the shoulder-joint.
This portion is divided into two heads, the aperture
of the division being a wide ol)tuse angle, with its
apex opposite the humeral head. This bifurcation
gives rise to the long and short heads of the hicej^s,
the former being inserted into the outer aspect of the
head of the coracoid just beyond the glenoid cavity,
while the short head is attached to the distal angle
of the ulnar tuberosity of the humerus (Figs. 24,
28, 29, and 32), This flat, proximal tendon extends
somewhat beyond the bifurcation, but soon is usurped
by the commencement of the carneous portion of the
muscle, which is correspondingly broad and subcom-
pressed. After forming the muscular belly, the fibres
as they pass down rapidly converge to an apex, and
finally form the strong cord-like tendon for insertion,
which passes to the ulna and is attached to that
bone, immediately in front of the articular cavity
for the trochlear surfaces of the distal extremity of the
humerus.

The biceps is a powerful flexor of the forearm, and
this is its chief function.

70. TJie deltoid ^ is partially divided into two
portions ; one long narrow head arises tendinous from

^ A very excellent account of the deltoid muscle in birds is also
given us by Gadow (loc. cit., p. 230), together with a synonynay
which is herewith republished . —

118 THE MYOLOGY OF THE RAVEN.

the clavicular process of the scapula and from the
contiguous surface of the clavicle itself (Fig. 33).
These fibres wind I'ound the Ijack of the shoulder-
joint, and are joined l)y the fibres that arise upon the
entire outer surface of the large os humero-scapulare.
These latter are inserted upon an extensive area upon
the ancoual aspect of the bone, beyond the humeral
articular head, and down nearly the entire length of
the shaft below it, making room for the latissimus dorsi
at the point where it is inserted. The first-mentioned
division of the muscle does not enter apparently into
this part of the insertion, Imt becomes more intimately
blended with the bulk of the muscle as the rapidly con-
verging fibres proceed down the humeral shaft, until
both blend to form a strong and distinct subcylindrical
tendon, which is inserted into the proximal side of the
tubercle of the external condyle, and aliove the insertion
of the extensor nietaeaypi radialis longus muscle of the
antibrachium.

*' 69. M. DELTOIDEUS MAJOU.

Le grand releveur de Vhumerus. Vicq d'Azyr.
Acliselheber. Merrem, p. 154, No. 14. ,

Le deltoide. Cuvier.

Aeusserer Oberarmstrecker. Wiedemann.
Deltoides. Meckel, System, p. 310, No. 1.
Deltoideus major -\- minor. Tiedemann, §§ 253, 254.
Deltoideus majm\ Riidinger ; Heusinger, p. 183.
Selenka, p. 116, No. 42.

„ „ De Man, p. 107 ; Carlsson, p. 20.

,, ,, Fiii-biinger, Mtyrph, Jahrh., xi.

Deltoideus sujjerior. Schopss, p. 117, No. 18.
Grosser deltaformiger Muskel. Prechtl, § 36.
Deltoid. Forbes, Tuhina^xs, p. 30.

Deltoideus s. axillaris. Flirbringer, Morph. JaJirh., v.
Deltoide 2)osti-rieur. Gervais et Alix, p. 23.
Sous-eirineux et deltoide posterieur. Alix, p. 394-96.
Tensor patagii lowjus {superficialis part.). Watson, p. 88."

THE MUSCLES OF THE UPPER EXTREMITY. 119

For nearly its entire lengtli the deltoid is intimately
attached to the triceps by fascia, more especially to its
long head, where it comes in contact with that sli23 of
the former which passes round the shonlder-joint.

71. The tricej)s^ muscle in the Eaven is divided into
three portions, an internal and external head, and a long
or scapular head, but so loosely attached is the latter to
the remaining two heads of the triceps, that it almost
amounts to two muscles in this bird.

The long or scapular head arises, somewhat tendinous,

^ Agreeing with what I found in my dissections upon Corvus,
three years ago, Gadow has also declared that " Der Triceps
brachialis besteht bei den meisten Vcigeln aus di'ei in der Regel von
einander getrennt bleibenden Theilen, namlich eine P. scapuli-
cubitalis und 2 P. humero-cubitales (ext. et intern.) " [loc. cit., pp.
263, 264). He gives the following synonymy, to wit : —

"81. M. TRICEPS CUBITI S. ANCONiEUS.

Le grand extenseur du coude. Vicq d'Azyr, p. 571.

Extenseur de Vavant bras. Cuvier.

Ohne Namen. Merrem, p. 155, No. 1 und 2.

Lange ausserer und innerer Ellenbogenmuskel. Wiedemann, pp.

86-89.
Anconceus longus, Irevis et brevissimus. Tiedemann, §§ 264, 265

266.
Der lange, kurze und kleinste Ellenbogenknorrenmuskel {An-

conceus longus, brevis et quartus). Heu singer, p. 185, No.

16, 17,18.
Strecker des Yordearmes. Meckel, p. 331, No. 9.

Schupss, p. 190, No. 23.
Langer, kiirzerer, und kleinster Strecker. Prechtl, §§ 43, 44, 46.
Triceps extevisor cubiti et anconeus. Eeid, p. 142.
Triceps bracliii. Selenka, p. 126, No. 53.
De Man, No. 19.
„ ,, Forbes {Tubinares), p. 30 ; Carlsson, p. 21.

Triceps extensor cubiti. Watson, p. 90.
La longue p)ortion du tricejjs brachial, le vaste externe et le vaste

interne. Gervais et Alix, pp. 25 und 26.
Triceps brachial. Alix, p. 404.
Anconeus {sca2)ularis -\- coracoideAis). FxivhringeY, Morph.Jahrb., xi."

120 THE MYOLOGY OF THE RAVEX.

from tlic superior aspect of the scapula, upon a circum-
scribed area just posterior to the glenoid cavity (Fig. 24);
the fibres form a ratlier thick, flattened muscle, which,
winding round the shoulder-joint, beneath the deltoid,
and parallel to the fibres of the scapular head of that
muscle, becomes feebly blended with those of the external
head of the triceps, as they together pass down on the
postero-lateral aspect of the humeral shaft. The con-
nection of this part of the muscle along its course, in
this locality, with the other heads of the triceps, is
most intimate through the closely binding fascia, rather
than through any blending that takes place among the
fibres of the several portions.

Just above the elbow, the fibres of the scapular head
of the triceps converge and merge into a strong cord-
like though flattened tendon, which passes to the radial
side of the olecranon process of the ulna to become
inserted there, near its base.

The internal and extSrnal heads of the triceps are
intimately blended except at their proximal extremities.
In this last situation they form a well-marked bifurca-
tion, where each arises from the anconal surface of the
shaft of the humerus ; the internal head towards the
deltoid attachment, and the external head running into
the pneumatic fossa. Between the limbs of the mus-
cular bifurcation thus formed arises the supraspinatus
muscle. The external head of the triceps in this locality
also of itself bifurcates in order to allow the teres et
infraspinatus muscle to attach itself to the humeral
shaft (Fig. 27).

The internal and external heads of the triceps become
blended in their attachment down the anconal aspect of
the bone to a point below the insertion of the deltoid.
Here they form a flat tendon, partly muscular, which is
quite broad, and that passes over the elbow-joint to

THE MUSCLES OF THE UPPER EXTREMITY.

121

insert itself into the entire under surface of the ole-
cranon process of the ulna.

The triceps is the great extensor of the antibrachium,
and direct antao-onist of the hrachialis anticus and
biceps. Its scapula or long head also materially pro-
tects that part of the shoulder-joint over which it passes,

tcruZ^yrv of ext:.irv&t. rtLciy.Zon^vor.
jfrona^T' long vis.

Fig. 36. — Bones of the forearm and liaiid of a Raven ; palmar aspect, and slightly-
dislodged from their normal positions in order the better to show the
origins and insertions of muscles for which the drawing is designed.
Life-size, by the author, from a specimen in his own collection.

and assists other muscles of that region in retaining the
humerus in its shallow socket.

72. The hrachialis atiticus is a very well-defined and
neat little muscle in the Raven (Figs. 24, 29, and 37). It
arises almost entirely fleshy from a circumscribed area on
the inner side of the anconal aspect of the distal extremity

122 THE MYOLOar OF THE RAVEN.

of the humerus, its fibres passing directly over the elbow-
joint on that side to insert themselves upon the hiteral
surface of the proximal end of the ulna quite up to
the margin of the greater sigmoid cavity.

This muscle protects the anterior aspect of the elbow-
joint, and assists the hicej)s in flexing the antibrachium
upon the brachium.^

MUSCLES OF THE FOREARM AND HAND.

As we pass towards the hand in a bird, and this Raven
forms no exception, we find the muscles deviating still more
from the musculature of the corresponding parts among
the Mammalia. Muscles arise and tendons depart from
their more common tracks to be inserted as best meets
the main function of this limb in Aves — its modification
to an instrument for the purposes of flight. Many of
these muscles I can make out, but quite a number of
them are not noticed by Owen, Huxley, Mivart, Garrod,

^ Flirbringer describes this muscle as the brachialis inferior, and
in this he was followed by Gadow, who retained the same name for
it, and in Bronn's Klassen presents the following synonymy {Joe.
ciL, p. 262) :—

" 80. M. BKACHIALIS INFERIOR.

Le court JlecJdsseur de Vavant bras. Vicq d'Azyr, p. 572.

,, ,, ,, ,, ,, ,, Gervais et Alix, p. 27.

Ellenbogenbeuger [Flexor parvus vlno'). Wiedemann, j). 89.
Brachialis intermis. Tiedemann, p. 312, No. 2.
d'Alton, p. 25, No. 23.
,, ,, Rudinger, p. 100; Heusinger, p. 184, No. 11.

Selenka, p. 125, No. 52.
De Man, No. 18.
,, ,, Watson, p. 91.

„ ,, Carlsson, p. 27.

Kleiner Beuger. Meckel, p. 325, No. 2.
Kurzer Beuger des Vorderarmes. Schopss, p. 141, No. 28 {Flexor

hrevis antihrachii).
Innerer Armmuskel (Einzieher des Vorderarmes). Prechtl, § 47.
Bracldalis inferior , Fiirbringe)'."

THE MUSCLES OF THE UPPER EXTREMITY. 123

or Forbes, and in the absence of tbe works of other
authors I shall be obliged to name tliern as best I may,
and as the functions they perform seem to indicate.

I must believe that if Mr, Garrod had paid as much
attention to certain groups of the muscles of the arm,
forearm, and j^inion in birds, as he did to certain re-
stricted groups of muscles of the lower extremity, he
would have discovered characters of value in classifica-
tion fully as significant as those he so ably elucidated in
the latter region. His excellent observations upon the
methods of origin and insertion of the tensor 2)''ftagii
hrevis point most emphatically to that fact. As I have
so often repeated elsewhere, we may say as we please,
but the classification of animals will only be placed
beyond all doubt when their entire morphology is hnoum
and correcthj comprehended, and duly utilized.

In the freshly plucked wing of a Raven we notice
how very prominent the muscles of the forearm and
pinion are, more particularly the tendons and muscles
upon the inner aspect of the forearm. In this locality,
it will be seen, as in the case of the tendon of the ex-
tensor metacarpji radialis longior, that they are almost
completely ensheathed in the common integuments, so
prominently do they stand out. In removing the skin,
we discover a strong fibro-elastic cord, which passes from
the under side of the proximal end of the ulna, and
joins each and every quill of the row of large feathers
of the wing, at a distance of about a centimetre or
less, all the way to the distal apex of the pinion, towards
which it gradually converges, and where it is finally
attached. This semitendinous cord plays the part, to
a certain deg-ree, of an antao-onistic tendon to the
tendon of the tensor patagii longus, it being by no
means an inefticient fiexor to the pinion upon the fore-
arm, and in closing the wing draws the hand towards the

124 THE MYOLOGY OK THE RAYEN.

antibrachium. This tensor of the posterior patagium,
of course, is covered by the folding of that membrane,
much in the same way as the tendon of the tensor
patagii longus is.

The forearm of the Raven presents us with the
following muscles for our examination, viz. : —

73. The extensor metacarpi 80. The anconeus.

radialis longior. 8J, The extensor indicis longus.

74. The extensor digitorum 8la. The flexor digitorum sub-

communis. limis.

75. The supinator brevis. 82. The flexor digitorum pro-

76. The flexor metacarpi radialis. fundus.

77. The pronator brevis. 83. The flexor carpi ulnaris.

78. The pronator longus. 84. The flexor carpi ulnaris

79. The extensor ossis metacarpi brevior.

pollicis.

73. The extensor tnetaearpi radkdis longior^ (or
longus as it is occasionally written) is one of the, if not

^ This important muscle Prof, (iadovv has termed the extensor
metacarpi ulnaris (which is probalily a misprint, and should be
radialis). He gives its synonymy (loc. cit., pp. 274, 275) as
follows : —

" 88. M. EXTENSOR METACARPI ULNARIS [rADIALIS 1].

Le long radial. Vicq d'Azyr, 1773, p. 575, No. 1.
Le radial. Cuvier.

Hinterer ausserer Handspanner. Merrem, p. 156.
Mittelhandstrecker. Wiedemann, p. 90.
Extensor metacarpi radialis longus. Tiedemann, § 268.
(Langer Speichen-Mittelhandstrecker.) Heusinger, p. 187, No.

22 +"'No. 21.

„ „ „ Schcipss, 145, No. 31.

.Extensor metacarpi i-adialis longtis. Selenka, p. 130, No. 59.
(Langer Speichen-Mittelhandstrecker.) Carlsson, p. 22.
„ „ De Man, No. 23.

„ „ „ Watson, p. 94.

Langer Speichenstrecker. Meckel, p. 333, No. 1.
Langer Mittelhandstrecker. Prechtl, § 52.
Extensor metacarpi longus. d'Alton, p. 27, No. 28.

THE MUSCLES OF THE UPPER EXTREMITY. 125

the, principal extensor of the hand upon the forearna.
It arises by two strong tendinous heads : the outer one
from the tubercle of the external condyle of the humerus,
just above the origin of the tendon of the tensor patagii
brevis ; and the inner and stronger one from a tubercle
found above the oblique trochlear facet of the distal end
of the same bone, for the radius. The median nerve
passes between the two heads, after which they unite to
form a large fusiform muscle, the highest of the group,
seen on lateral aspect of the arm, which, running
parallel with the ulna, becomes converted into a flat,
broad, and strong . tendon at about the middle of the
forearm. This passes directly on to become inserted
into the apex of the anchylosed first metacarpal of the
carpo-metacarpus.

The attachment that the tensor pataijii hrevis makes
with the present muscle has already been described
above. The distal extremity of the radius is distinctly
grooved in a lonoitudinal direction to admit of the
lodgment of the tendon of the extensor metacariyi
radialis longior, before it makes its final attachment.
Owen says that this muscle " raises the hand, draws it
forward toward the radial margin of the forearm, and
retains it in the same plane. In the Penguin this
muscle is extremely feeble, and the tendon is lost in
that of the tensor plicce alaris."

74. The extensor digitorum communis ^ is a smaller
muscle than the one just described, and arises from the

Le long supinateur. Gervais et Alix, p. 26.
„ „ „ Alix, p. 408.

Extensor carpi radialis. Selenka, p. 129, No. 58; Carlsson, p. 22.
Extensor carpi radialis Irevis. Selenka, p. 130, No. 59a."

^ This is 92. M. extensor digitorum communis of Gadow (Bronn's
Klassen des Thier-Eeichs, vi. Band, p. 282) and of Selenka (loc. cit.,
p. 131, No. 61).

126

THE MYOLOGY OF THE RAVEN.

liiimerus jminciliately below the tul)ercle of the external
condyle. It soon forms a, spindle-shaped muscle which
blends with the supinator hrevis as it passes it, and
terminates in a strong, cord-like, glistening tendon at
about the middle of the forearm. This tendon passes

Eixtan^ar"
OSS is mjeXa>-

vC^upuuitbi'

Fig. 37. — Radial aspect of left
ulna and radius of an
adult Haven, designed
to show origin and
insertion of muscles
of the forearm. Life-
size, by the author,
IVom the siiecimen.

through a groove in common with the tendons of the
JiexoT carpi radialis and the long extensor on the
palmar side of the distal extremity of the ulna, and in
passing down the front of the hand, first throws off a
small tendon which is inserted on the outer side of the

THE MUSCLES OF THE UPPER EXTREMITY. 127

base of the poUex digit ; after wliicli, it continues down
the anterior aspect of the shaft of the carpo-metacarpus
in a well-marked and somewhat oblique groove intended
for it, as far as the upper end of the proximal phalanx
of the middle digit. Here it lies beneath the tendons
of the other muscles, when turning to the median line
through a fibro-cartilaginous pulley intended for it, it is
finally inserted at a mid-point upon the anterior rim
of the proximal phalanx of the middle finger. Upon
lateral aspect of the outer side of the muscles of the
forearm, the extensor digitoruni communis lies next
below the extensor metacarpi radialis long lor, occupy-
ing the middle of the group there found.

75. The supinator hrevis^ is exposed after remov-
ing the last-described muscle, to which it is rather
closely attached by a tendinous connection. It arises
from the external condyle of the humerus, below the
origin of the tendon of the extensor digitorum com-
munis, and passing directly in a, nearly straight line to
the radius it becomes attached to the outer side of the
shaft of that bone for nearly one-third of its length.
This muscle, here quite feebly represented, acts, as in

^ " 84. M. ECTEPICONDYLO-RADIALIS.

Le court swpinateur. Vicq d'Azyr, p. 573.
„ „ „ Cuvier.

,, „ „ Gervais et Alix, p. 2G ; Alix, p. 407.

Anleger des Yorderarmes. Merrem, p. 155.
Aeusserer Speichenbeuger. Wiedemann, p. 90.
Suinnator. Tiedemann, § 262.

„ Schopss, p. 140, No. 27.

Supinator brevis. Heusinger, p. 185, No. 14.

Selenka, p. 12'J, No. 57 ; De Man, No. 22.
„ ,, Watson, p. 93 ; Carlsson, p. 23.

Riickwartswender. Meckel, p. 330, No. 8 und No. 7.
Antagonist der Niederzieher. Prechtl, § 51." (Gadow in Bronn's
Klassen, lac. cit., p. 269.)

128 THE MYOLOGY OF THE RAVEN.

the limbs of other Vertebrata, as a supinator, and feeble
antagonist to the powerful pronators to be described
further on.

76. The flexor metacaiyi radialis^ is the lowest of
the group of three muscles on the outer aspect of a
Raven's forearm. Its mode of origin is very interesting,
for we find it arises by two distinct tendinous heads,
the longer one coming off from the external condyle of
the humerus, where it overlaps the origin of the siqnna-
tor hrevis, while the shorter one arises from the ulna
just without and rather beyond the base of the olecranon
process. In common with the others of the group of
muscles to which it belongs, it has a fusiform shaj^e for
the proximal moiety of the forearm, and is intimately
attached for its middle third to the interosseous
membrane.

Rather beyond the middle of the shaft of the ulna the

^ Tiedemann used this name for the muscle here described, as will
be noted from Gadow's synonymy [loc. cit., p. 276) : — •

" 89. M. EXTENSOR METACARPl ULNARIS.

Le long flechisseu'i' du metacarpe. Vicq d'Azyr, p. 575, No. 5.
Cubital interne. Cuvier.

Hinterer innerer Handspanner. Merrem, p. 156 (?).
Oberer oder langer Mittelhandbeuger. Wiedemann, p. 91.
Flexor metacarpi radialis. Tiedemann, § 271.

,, ,, „ Heusinger, p. 188, No. 25.

Aeusserer Ellenbogenmuskel. Meckel, p. 335, No. 3.
Abductor metacarpi. Schopss, p. 150, No. 33.
Langer Mittelhandbeuger. Prechtl, § 55.
Extensor carpi ulnaris. Selenka, p. 131, No. 60.

,, „ „ De Man, No. 24 ; Watson, p. 95 ; Carlsson,

p. 23.
Le cubital posterietir. Gervais et Alix, p. 27.
„ „ „ Alix, p. 409."

[This muscle flexes the hand on the antibrachium, although it
arises from the external condyle. — R. W. S.]

THE MUSCLES OF THE UPPER EXTREMITY. 129

flexor metacarpi radialis tapers to a j)oint, then merges
into a strong, cord-like tendon, which passing through
the fibrous sheath to the outer side of the distal end of
the ulna, in common with the tendons of other muscles
previously alluded to, it goes directly to the prominent
process which juts from the proximal third of the
posterior aspect of the shaft of mid-metacarpal, where it
is firmly attached. This muscle, when brought fully
into action, is a powerful flexor of the hand upon the
antibrachium.

In the Ostrich this muscle arises from the lower third
of the ulna, while in the Penguin it is entirely absent
(Owen).

In Fig. 36 I have drawn the line occupied by the
tendons of this muscle, as well as the line of its axis,
which is supposed to be in the same straight line with
its tendon of insertion.

77. The p)'onator hrevis^ is the uppermost of the

^ Of the pronator group of mviscles in birds, Gadow records the
following synonymy {loc. cit., pp. 266, 267) : —

" 82. Mm. entepicondylo-antibrachiales.

" A. — ^Entepicondylo-eadiales.

" /. Pronator suhlirais s. hrevis.

Le viuscle q2ii tient lieu du -pronateur rond. Vicq d'Azyr.

Kurzer Speichenbeuger. Wiedemann, p. 89.

Pronator p7'imus s. hrevis. Tiedemann, § 260.

Beuger des Yorderarms. Meckel, p. 326.

Pronator brevis. Schopss, p. 137, No. 25 ; d'Alton, p. 26 ; Riidinger,

p. 113 ; Selenka, p. 127, No. 54 ; Carlsson, p. 22.
Kurzer Niederzieher des Vorderarms. Prechtl, § 49.
Pronator sublimis. De Man, p. 112.
Pronator teres. Gervais et Alix ; Watson, p. 102.

" //. Pronator profundus s. longus.
Le radial externe. Vicq d'Azyr.
Langer Speichenbeuger. Wiedemann, p. 89.

K

130 THE MYOLOGY OF THE RAVEN.

pair of powerful pronators in the forearm of this bird.
It arises by a tendon rather above the internal condyle
of the humerus ; the fibres, forming a thick fusiform
muscle, pass obliquely across the interosseous space to
become inserted on the ulnar side of the shaft of the
radius, just beyond the junction of the proximal and
middle thirds. This muscle is somewhat compressed

Pronator secundus s. longus. Tiedemann, § 261 ; Rudinger, p. 113.

Beuger des Yorderarms. Meckel, p. 326.

Pronator longus. d' Alton, p. 26 ; Schopsp, p. 138, No. 26 ; Selenka,

p. 128, No. 55.
Langer Niedeizieher. Prechtl, § 50.
Rond pronateur frofond. Alix.
Pronator jyr of undus. De Man, p. 112.

" B. — Entepicondylo-ulnaris.

Flexor hrevis ulnae. Wiedemann, p. 93.

,, ,, ,, Tiedemann, ^ 263 ; Heusinger, p. 185, No. 15.

Ohne Namen. Meckel, p. 328, No. 5.
Flexor lyrofundus interior gallhtaceorimi. Schopss, p. 144, No. 30.

,, „ „ „ Selenka, p. 128, No. 56.

Kurzer Beuger der Elle. Prechtl, § 48.
Ancone interne. Alix, p. 408."

This synonymy is immediately followed by a concise description
(in the same work) of the proaators, they agreeing substantially
with my 2jronator hrevis and longus (Nos. 77 and 78) respectively.
Then comes a description of Prof. Gadow's m. entepicondylo-ulnaris,
a muscle, according to his description, that takes origin from the
internal condyle of the humerus and is insei'ted into the ulna. Thus
far I have failed to discover the particular muscle to which he
refers. From the synonymy given above it will be observed that
Tiedemann and others designated it as the Flexor hrevis idnce. Prof.
Gadow's descriptions are in the main very brief, and the figures of
his plates (for the most part copied from Alix, De Man, and
Watson) are by no means satisfactory, so that it is possible, or more
than likely, that one runs the chance of mistaking the muscle he
refers to in some cases. I have omitted the synonymy in several
instances, as will be noted below.

K 2

132 THE MYOLOGY OF THE RAYEN.

from above downwards, and its tendon of origin is seen
to spread out over its proximal moiety on either side.
In addition to its being a powerful muscle of pronation,
it may act also as a flexor of the forearm upon the
arm.

78. The 'pronator longus^ i^ even a stronger muscle
than the preceding, and possesses considerably more
bulk. It comes off by a strong tendon from the middle
of the internal condyle of the humerus, its fibres
forming a massive ellipsoidal muscle, which, passing
parallel to the pronator hrevis, is inserted into the shaft
of the radius, just beneath that muscle and rather pos-
terior to it. The jyronator longus is in intimate relation
with the hrachialis a^iticus, the flexor digitorimi 2?ro-
fundus, and other muscles lying beneath it. Its action
is much the same as we described for the short pronator
in the last paragraph.

79. The extensor ossis 7netacarpipolUeis'^i^,hyii\\

^ See footnotes under No. 77, antea.

2 I fail to find this muscle exactly described by Gadow as I have
it here, and, as I say under its description above, it may be the ext.
vietacarjji radialis brev. of Tiedemann, Watson, and others.

However this may be, Gadow, I see, desciibes the following (Joe.
cit., pp. 283, 284) :—

" 93. M. EXTENSOR POLLICIS LONGUS.

Le radial grele. Vicq d'Azyr, 1773, p. 574, No. 2.
Vorderer Handanleger. Merrem, p. 157, No. 4.
Hiilfsmuskel des Mittelhandstreckers. Wiedemann (1).
Extensor metacarjn radialis hrevis. Tiedemann, § 269.

,, ,, ,, „ Heusinger, ]). 187, No. 23.

,, ,, ,, ,, Schtipss, p. 148, No. 32.

„ ,, „ ,, Watson, p. 95.

Kurzer Speichenstrecker. Meckel, p. 334, No. 2.
Extensor pollicis longus. Elidinger, p. 131.

,, ,, „ Selenka, p. 133, No. 63.

„ „ „ De Man, No. 27, Carlsson, p. 23.

THE MUSCLES OF THE UPPER EXTREMITY. 133

odds, the smallest and most delicate muscle of the fore-
arm in our present subject. It arises immediately in
front of the "greater sigmoid cavity" of the ulna, in
intimate relation with the fibres of the anconeus, and
from the interosseous membrane close to the head of the
radius, and I believe it may even extend to that bone
for attachment. Its fibres form a very delicate, straight
little muscle, completely hidden from sight by the
others that surround it, and which does not exceed
three centimetres in length, when it becomes converted
into a tendon of absolutely hair-like dimensions. This
passes directly to the wrist, parallel and in close connec-
tion with the extensor nietacarpi radialis longior, to
be inserted with it at the base of the first metacarpal,
to its palmar side. In its action this little muscle plays
the part of a feeble extender of the hand upon the
forearm.

Mivart says that this " muscle may be double and
very voluminous, as e.g. in the Chameleon. Even in
very near allies of man (the Anthropoid Apes) it ends
in two tendons, one going to the trapezium, the other to
the metacarpal of the thumb" (Ele^n. Anat., p. 336).

Kurzer Mittelhandstrecker. Prechtl, § 53.

Abductor du pouce. Gervais et Alix, p. 409 ; Alix, p. 409.

" Dieser Muskel entspringt, bedeckt vom Ext. indicis, von den
einander zugekehrten Fliichen des Radius und der Ulna, und zwar
vom proximalen Theile derselben. Hierdurch erhalt der Mnskel
eigentlich zwei mehr oder weniger getrennte Ursprungskopfe.
Dieselben vereinigen sich zu einer Sebne, die erst die Aussen — oder
Vorderfliiche des Radius begleitet und dann auf der dorsalen oder
Extensorseite von eineru eigenen Bande gebalten, iiber das Os carpi
radiale lauft, um sich schliesslich an dem Vorsprucge des Meta-
carpale I. zu inseriren

^^'BeiParadisea, Oriolus, Corxusn&ch De Man ebenfalls zweikopfig
und zwar ist der radiale kopf der stiirkere."

134 THE MYOLOGY OF THE KAVEN.

It is also found in tlie Horse, where the thuml) is
entirely absent. I fail to find an extensor metacarpi
radialis hrevior, so frequently described for birds by
other authors, and it is just possible that this is the
muscle alluded to by them. Of it, Owen says that it
arises below the preceding [e.xtensor 7netacarpi radialis
lon(jior~\ from the uhiar edge of the radius, and is inserted
into the phalanx of the thumb immediately beyond the
tendon of the preceding muscle [extensor metacarpi
radialis longior']. The two tendons are quite distinct
from one another in the birds of prey, the Ostrich and
Parrots, but unite at the lower end of the forearm in
the Anatidce, Phasianidce, and Gruidce'' (Anat. of Verts.,
vol ii. pp. 98-99).

80. The anconeus'^ is a very powerfully developed
muscle in the forearm of the Raven. It arises by a
short, though strong, subcylindrical tendon from the
lower aud back part of the external condyle of the
humerus, and passes directly to the latero-radial side of
the shaft of the ulna, alomx which it attaches itself to a
point somewhat beyond its middle (Figs. 30 and 37).

On its inner side it is in intimate relation with the

1 " 83. M. ECTEPICONDYLO-ULNARIS.

Lejlechisseur in-ofond de Vavant bras. Vicq d'Azyr, 1773, p. 573,

No. 8 ; Cuvier.
Oline Namen. Merrem, Fig. 3, v.
Kurzei* EUenbogenstrecker. Wiedemann, p. Ul.
Unterer Kurzer oder vierter EUenbogenstrecker. Meckel, p. 329,

No. 6.
Flexor antibrachii ]y)-qfundus. Schopss, p. 142, No. 29.

Gurlt, p. 23, No. 6.
„ ,, ,, Watson, p. 62.

Ancone. Alix, p. 407.
Anconceus quartus. De Man, p. 115. (Von Tiedemaun, Prechtl,

und Selenka nicht erwiihnt.) " (Gadow, loc. cit., p. 268.)

THE MUSCLES (.)F THE UPPER EXTREMITY. 135

deep flexors of the forearm, and in that region its
investing fascia shows a strong disposition to become
tendinous, as may be observed from its striated and
glistening white coh)ur.

81. The extensor indicis long us} — li\ Cor vus, onr

^ Gadow makes the following remarks, and presents the following
synonymy of this muscle, to wit : —

"94. M. EXTENSOR INDICTS LONGUS.

L'extenseur externe du doigt. Vicq d'Azyr, 1773, p. 574, No. 3.
Strecker des ersten und zweiteu Gliedes des zweiten Fingers.

Tiedemann, § 278.
Aeusserer oder hinterer Strecker des ersten uud zweiten Gliedes

des zweiten Fingers + Hulfsmuskel des iiasseren Streckers.

Heusinger, p. 193, No. 35 ; p. 194 und p. 196, No. 36.
Eigener Strecker des zweiten Fingers. Meckel, p. 344, No. 2.
Extensor indicis in'ojjrhis longns. Schopss, p. 159, No. 38.
Strecker des grossen Fingers. Prechtl, § 58.
Extensor digiti indicis proj^rius (longus et hrevis). Selenka, p. 132,

No. 62 ; Watson, p. 97 ; Carlsson, p. 24.
Extensor indicis longus. De Man, No. 27.
Extenseur de la deuxieme iTihalmige du doigt median. Gervais et Alix,

p. 28.
Extenseur de la deuxieme fhalange du deuxieme doigt. Alix, p.

414.

" Der besondere Strecker des zweiten Fingers besteht meistens
aus zwei nur mit ihren Endsehnen vereinigten Muskeln.

" I. Der grossere, stets vorhandene Theil liegt dorsal in dem von
Ulna und Radius eingeschlossenen Raume und entspringt fleischig
gewohnlich von den mittleren zwei Dritteln der dorsal- und ulnar-
warts schauenden Fliiche des Radius, kann aber auch Fasern von
der gegeniiber liegenden Flache des proximal en Endes der Ulna
erhalten.

" Der Muskel geht am letzten Drittel des Vorderarmes in eine
Sehne liber, welche liber eine Rinne auf der oberen Flache des
distulen Endes der Ulna lauft ; sie geht dann dorsalwtirts Uber
das Metac. II. bin zur Radialseite des zweiten Fingers, ist durch
Bander an das Gelenk des ersten Gliedes niedergedrlickt und
inserirt sich an der Radialseite des Caput, phal. I. und der Basis
phal. II. dig. II. Ist wie bei Lamelliroslres, Grus, Numenius,

136 THE MYOLOGY OF THE EAVEN.

present suljject, it arises from the iiifero-ulnar aspect of
rather more than half of the proximal moiety of the
shaft of the radius, as a small and quite inconspicuous
strip of muscle. This becomes converted into a long-
tendon, stronger than we might be led to expect from
the size of the carneous portion of the muscle, which,
passing over the groove at the distal end of the ulna, in
company with the tendons of other muscles already
described above, passes directly down on the anterior
aspect of the hand to be inclosed in a fibrous sheath in
front of the superior aspect of the first phalanx of the
index digit ; passing which, it is finally inserted into
the anterior and upper rim of the distal phalanx of the
same finger.

Struthio ein drittes Fingerglied vorhanden, so erstreckt sich die
Sehne bis an die Basis desselben.

" Auf dem Metacarpal-Phalanx-Gelenke findet sich oft ein Sesam-
bein. In der Carpalgegend wird die Sehne von der des Ext. dig.
comm. bedeckt, spaterhin aber wird sie zur oberflachlichsten und
bedeckt ihrerseits die Zeigefingersehne des erwjihnten Muskels.
Der Ursprung des Muskels dehnt sich auf nahezu die ganze Lange
des Radius aus bei Cormis und Paradisea ; bei den Raptores,
Columbce, Rasores, Grallce entspringt er ungefahr von der mittleren
Halfte, d. h. er lasst die Enden frei ; er konimt nicht vom
proximalen Drittel, geht dafiir aber bis an das distale Ende bei
Psittacus. Bei den Spheniscidce ist er sehr schwach.

" II. Der zweite, klirzere Kopf entspringt vom Dorsalrande
entweder des distalen Endes des Radius, vom Os carpi radiale oder
von der Basis des Metac. II. ; seine Sehne verbindet sich in der
Nahe des distalen Endes des Metacarpus mit der Endsehne des
griisseren Theiles.

" Dieser Hlilfsmuskel findet sich nicht bei Corviis, Paradisea,
Fulica, fSpheiiiscus.

" Vergleichung. Entspringt nur unvollkommen dem Ext. indicis
proprius des Menschen, da dev Mnskel bei den Vugeln fast
ausschliesslich vom Radius anstatt von der Ulna entspringt."
(Bronn's Klassen, loc. cit., pp. 285, 286.)

THE MUSCLES OF THE UPPER EXTREMITY. 137

81 A. The Jiexor digitorum suhlmiis. — In dissecting
the muscles of the forearm of a Raven one meets with
a tendinous, somewhat muscular band, closely adherent
to the integument, that stretches from the internal con-
dyle of the humerus to the wrist, and from this latter
point sends down a tendinous cord into the hand. I
have not shown this structure in mv figures, although it
is generally recognized among the muscles by ornitho-
tomical writers. Gadow describes it as follows : —

"90. M. FLEXOR DIGITORUM SUBLIMIS.

L'extenseur grele de la ^^ar^/e qui tieiit lieu de doigt. Vicq d'Azyr,

p. 572.
L'adducteur de la j^Temiere 2>halanye. Cuvier.
Fingerspanner ('?). Merrem, p. 157.

Oberer oder langer Mittelhandbeuger. Wiedemann, p. 91.
Vorderer Strecker des ersten und zweilen Gliede^ des zweiten

Fingers. Heusinger, p. 191, No. 32.
Oberflacblielier langer Fingerbeuger. Meckel, p. 346, No. 3.
Flexor digitwum superjicialis. Schopss, 161, No. 39.
Flexor digitorum sublimis. Selenka, p. 135, No. 66.

De Mau, No. 31.
„ ,, „ Watson, J). 99.

,, ,, ,, Carlsson, p. 27.

Le 2)etit ^ja^?Mai?'e und Jl,ecJtisseur de la 2»'6inie)-e -pJialange du second

doigt. Gervais efc Alix, p. 29 ; Alix, p. 416.
(Tiedemann und Prechtl ervvahnen diesen Muskel nicht.)

" Der oberflachliche Beuger der Finger wird von einer starken
aponeurotischen Fascia bedeckt, welche nach Abtragung der Haut
auf der Volarflache des Unterarmes erscheint. Sie erstreckt sich
vom Cond. int. humeri langs der Yolarfliiche der Ulna zur Hand-
wurzel, wo sie sich mit einem Zipfel am proximalen Rande des Os
carpi ulnare ansetzt, wahrend der iibrige Theil volarwarts an
demselben Knochen sich befestigt. Der betreffende Muskel ent-
springt im allgemeinen von der dem Knochen zugekehrten Flache
dieser eigenthiimlichen, gewohnlich starken Sehne : die Insertions-
sehne des Muskels lost sich von letzterer etwas oberhalb der
Handwurzel ab, und dann etwas volarwarts in einer eigenen Rinne

138 THE MYOLOGY OF THE RAVEN.

iiber das Os carpi ulnare, dabei von Bandern in ihrei' Lage gehalteu
und darauf an der Mittelhand herab sich zur Eadial-Vorderflaclie
des Plial. I. dig. II. zu begeben.

"Der Muskel und das Humero-carpal-Baud sind grossen Ver-
Rcliiedenheiten vinterwoi-fen. 1st das Sebnenband sehr stark, wie
bei Anser, Carlo, Mormon, Uria, Tubinares, Heliornis, Otis, Oedicne-
mus, Tantalus, Parra, Raj)tores, Corvidce, so ist der Muskel schwach
und dlinn ; bisweilen wie bei Frocellaria und Spheuiscus gauz rudi-
mentar, die Endsehne bleibt aber gewohnlich erhalten und geht
zum Index. Ist dagegen das aponeuroti.sche Band schr schwacb ent-
wickelt, wei bei Gallus und Numida, so ist der Muskel auf dessen
letztes Drittel beschrjinkt und vereinigt sich bisweilen mit der
Sehne des Interosseus dorsalis.

" Der Ursprung der Muskels ist in verschiedener Ausdehnung
auf die tiefe Flacbe des Sehnenbandes bescbrankt bei den Schwimm-
Sumpf- und Tagraub-Yogeln, oder seine Fasern entspringen auch
von den sich an die gauze Lange der Ulna zwischen M. flex. dig.
profundus und M. flex, carpi ulnaris anheftendeu Theile der Aponeu-
rose {Buho, Podargus, Gajjrimulgus). Bei Parra chalcoptera war
das Sebnenband sehr stark, der Muskel schwach und ging nicht bis
an den Phalangen, sondern verier sich ain Os carpi ulnare und
etwas weiter abwarts Jiuf und in den dortigen Aponeurosen.

" Die Insertion wechselt ; jedoch scheinen die Yerschiedenheiten
von geringer Bedeutung zu sein. Am haufigsten (Pasores, Pa.ptores)
inserirt sich der Haupttheil der Endsehne an einer kleinen An-
schwellung auf der Radial vordertlache der Basis pbal. I. dig. II.,
wahrend der Rest sich schriig weiter distalwjirts auf derselben
Phalanx verliert. Zwischen diesen beiden Insertionstheilen tritt
die Endsehne des M. flex, profundus hindiirch, welche die von ihr
perforirte Sehne des M. flex sublimis schon im Bereiche des Meta-
carpus begleitet hatte.

" Bei Psittacus und Volumha setzt sich die Endsehne bis zur Basi.s
phal. II. fort. Fiir Otis giebt Schiipss Insertion an der Basis pbal. I.
an. — ■ Fine Insertion der Endsehne am Daumen scheint nicht
vorzukommen, wohl aber erstrecken sich Sebnenzweige des Humero
carpal-Baudes bis auf die Metacarpalia und den Daumen. Bei
Struthio fehlt der Muskel sowohl als auch ein zwischen Humerus,
Ulna und Carpus ausgespanntes Band.

" Vergleichung. Entspringt mit Modificationen dem M. flexor
digitoium sublimis s. perforatus der Siiuger und theilweise der

THE MUSCLES OF THE UPPER EXTREMITY. 139

oberflachlichen Beugemiiskiilatur der Reptilien und Amphibien. '
(Bronn's Klassendes Thier-Eeichs, vi. Band, pp. 278, 279.)

It is a significant fact that neither Tiedemann nor
Prechtl, two very close observing anatomists, described
this muscle, nor had anything to say about it in their
works.

82. Tlie Jiexor cUfjitorum profundus^ arises Heshy
by two strong heads from the proximal extremity of the
uhia. Between these heads the hrachialis anticus
muscle passes to be inserted upon the same bone.

The radial head of the Jiexor digitorum profundus
comes off immediately in front of the articular humeral
facets and the tendon of the biceps, at about the middle
point on the shaft, while the remaining head arises from
the under side of the shaft as far over as tlie olecranon
process. These heads, as the fibres pass wristwards,
soon merge with each other, and the common muscle
thus formed is attached for some little distance to the
shaft of the ulna beneath it. After that, it rapidly con-
verges to a point, and near the middle of the shaft
becomes converted into its tendon. This passes to the
ulnar side of the carpus through the various fascia-
sheaths and fibrous loops there found, and once more
emerges below the first metacarpal. From this point
the strong tendon makes directly for the anterior aspect
of the superior margin of the proximal phalanx of the
index digit. Here it is again confined in a fibrous

^ This muscle is designated also by this name in the works of
Selenka (Bronn's Klassen des Thier-Reichs, p. 136, No. 67), De Man,
Watson, and Carlsson ; likewise Gadow (foe. cit., p. 279) so defines
it for his muscle No. 91, presenting us with a synonymy of the same'
referring to the writers dating from Vicq d'Azyr, as well as with
a concise account bringing its history up to our present knowledge
of its anatomy in Aves.

THE MUSCLES OF THE UPPER EXTREMITY. 141

sheath, which it passes through to finally insert itself
into the ulnar side of the base of the distal joint of the
index fing-er.

o

Mivart says that "the^exo^' ^jro/'wwdiis digitorum in
man possesses an exceptional distinctness and subdivi-
sion. As has been said, it may be intimately united
with the sublimis, as it may also be (even in Monkeys)
Math the Jlexor longus poUicis. When distinct from the
latter, it may yet send a tendon to the thumb, as in
Nycticehus. It may end in but one, or at most two tendons,
as in birds. In the complete separation of this muscle
from the one next mentioned \_flexov longus pollicis]
man differs from all the Apes" [Elein. Anat., p. 332).

83. The jiexor carpi ulnaris^ is an exceedingly in-
teresting muscle, and is the one that makes up the
fleshy mass on the under side of the forearm for its
posterior moiety. On the ulnar side of the olecranon
process of the larger bone of the antibrachium we find a

1 " 85. M. FLKXOR CARPI ULNARIS S. ENTEPICONIJYLO-CARPALIS.

Le cubital interne. Vicq d'Azyr, 1773, p. 573, No. 6.

„ „ „ Cuvier.

Ausdehner des Arms + Kegierer der Armfedern. Merrem, p. 155,

No. 6 und 7.
Langer EUenbogenbeuger. Wiedemann, p. 22.
Flexor carjn tdiiaris. Tiedemann, § 272.

,, ,, „ Hevisinger, p. 189, No. 26.

,, ,, ,, Selenka, p. 133, No. 64.

,, ,, ,, Riidinger, p. 158.

„ ,, ,, Watson, p. 93 ; Carlsson, p. 29.

Innerer Ellenbogenmuskel. Meckel, p. 336, No. 6.
Langer Beuger der Handwurzel + M. rector remigum secundi

ordinis. Schopss, p. 154, No. 35, und p. 88, No. 4.
Antagonist des langer Mittelhandstreckers. Prechtl, p. 54.
Cubital anterieur. Gervais et Alix, p. 29.

Metacarpien palmaire interne. Alix, p. 412, pi. II., fig. 2, No. 15,
16, 17." (From Gadow, Bronn's Thier-Reichs, vi. Band, p. 270.)

142 THE MYOLOGY OF THE EAVEN.

fibro-cartilaginoiis loop developed ; one end of this is
attached to the middle point of the posterior aspect of
the internal condyle of the humerus, while its other end
is fast to the ulnar side of the base of the olecranon pro-
cess. We will call this the humero-ulnar pidlcy. Now,
the origin of the flexor carpi ulnaris, the muscle under
consideration, is concerned with this very structure, for
we find that the muscle arises by two strong tendons ;
the one on the side towards the olecranon process, pass-
ing through the humero-ulnar indleij, makes fast to the
back of the internal condyle of the humerus ; while the
outer tendon attaches itself to the same protuberance,
but at its outer aspect, and w^ithout the "pulley."

These tendons are somewhat flattened, and each about
a centimetre long, when tliey merge into a strong, fusi-
form muscle, occupying rather more than the posterior
half of the under side of the forearm. From the an-
terior apex of this muscle a powerful and subcylindrical
tendon stretches directly to the back of the nlnare ossi-
cle of the carpus, where it makes an extensive attach-
ment. Just before reaching this ossicle the tendon of
the flexor carpi ulnaris differentiates off a small tendi-
nous slip, wdiich, passing through a fibrous loop at the
ulnar side of the carpus, goes obliquely downwards to
the tendon of the flexor dlgitoriiin "profundus, and
merges with it at a point about opposite the distal apex
of the pollex digit ; or, more strictly speaking, is con-
tained in the same sheath with this tendon of the flexor
digitorum profundus tendon as far as the anterior rim
of the proximal phalanx of the index digit, where this
offshoot becomes inserted.

In its action this muscle is a powerful flexor of the
hand upon the forearm, and its tendinous offshoot at
the carpus is not an inefficient aid to the action of the

144 THE MYOLOGY OF THE RAYEX.

flexor dlgitorura j^rofundus. Owen calls this muscle the
flexor' metacarpi idnaris, and says of it in the Apteryx
that it " arises beneath the forearm from the internal
pulley of the ulna ; continues fleshy to the pinion ; and
is inserted, first into the ulnar carpal bone, then into the
ulnar phalanx. The latter insertion is wanting both in
the Ostrich and Penguin " {Anaf. Verts., vol, ii. p. 99).

When we come to examine it carefully in the Raven,
especially the distinctness of its external head, and a
median fascia that runs longitudinally through its belly,
and finally its distal tendinous ofishoot, it is not difficult
for us to imagine that the muscle originally was com-
posed of two parts, or perhaps two distinct muscles
(Figs. 31 and 36).

84. The flexor caipi idnaris hrevior^ is a muscle
that I do not find described in any work at present
available to me, and which I here provisionally bestow
this name upon in view of the fiict that the flexor carpi
idnaris is often split up into several portions in some
other vertebrates, and this is more than likely an in-
stance of it. And then, again, I was influenced in my
decision from the additional fact that in its action it
assumes in p>art the function of t\\(i flexor carpi idnaris
in those animals where it does occur normally. The
flexor carpi ulnaris hrevior in the Raven, however, not
only to a limited extent flexes the hand upon the fore-
arm, but by its peculiar carpal insertion rotates the hand
towards the side of the body — a very important action
durino; flioht.

This muscle arises from quite a broad area covering
the middle third of the upper side of the shaft of the

^ From Gadow's account (loc. cit., p. 272) I would say that this
muscle was represented by his Ko. 86. M. ulni-metacarpalis ven-
TRALis — the adductor metacarpi of Schcipss and of Watson.

cS a:

te a
5 o

'^ CD

'TIS

U(J THE MYOLOGY OF THE RAVEN.

ulna (Fig. 37), being found immediately beneath the
fiexor (Jujitorum profundus, or rather the anterior por-
tion of its muscular division, and the posterior third of
the tendon that comes off from it. The carneous portion
of the fiexor carpi ulnaris hrevior is entirely attached
to the ulna, while its exposed surface is covered by a
glistening, tendinous fascia. This merges, anteriorly,
into its true tendon, which passes from the apex of the
muscle directly to the carpus, well above the ulna's
shaft. At the carpus it winds around in front of the
radiale bone, from the ulnar side, to become inserted
upon the outer edge, near the top, of the anchylosed os
magnum of the carpo-metacarpus. This anterior portion
of the tendon is firmly held in its position by a strong
fibrous sheath, in which it plays.

This muscle can only be satisfactorily examined by
removing the other muscles of the forearm ; and, indeed,
I have taken up all the muscles of this division of the
pectoral limb in the order in which they most con-
veniently occur for examination. So that, for the
student to follow my work, he will find his task much
lightened by investigating them in the same order, i.e.
from 73 to 84 inclusive, as given in the list.

To complete our musculature of the upper extremity
we have yet to consider those muscles, several in num-
ber, which may be regarded as the intrinsic ones of
the hand. In the extensive chapter on the muscles of
birds, in his Anatomy of Vertebrates (vol. ii.), Profes-
sor Owen simply alludes to these muscles, and neither
gives their names nor otherwise describes them ; while
Professor Mivart has nothing to say about them so far
as they occur in Aves, in his Anatomy. Consequently,
we shall be oblio^ed to name them as their localities,
origins, and insertions seem to indicate.

THE MUSCLES OF THE UPPER EXTREMITY. 147

In the Raven they are found to be as follows : —

85. The extensor proprius pollicis. 89. The abductor minimi digiti.

86. The flexor brevis pollicis. 90. The flexor metacarpi brevis.

87. The flexor minimi digiti. 91. The interosseous dorsalis.

88. The flexor minimi digiti 92. The interosseous palmaris.

brevis.

85. TJie extensor proprius pollicis ^ is a neatly
arranged little muscle admirably adapted to the exten-
sion of the pollex digit. Spindle-shaped in form, it
arises from the ulnar side of the tendon of the extensor
metacai'pi radialis longior, and is inserted into the
antero-ulnar side of the pollex phalanx, being tendinous
both at its origin and its insertion.

A muscle of this name is described by human ana-

^ Selenka and De Man considered this to be an abductor of the
thumb^ and Gadow has given us the following synonymy for the
muscle : —

" 98. M. ABDUCTOR POLLICIS.

Innerer Daumenstrecker (Extensor jjollicis internvs). Wiedemann,

p. 93 ; Schopss, No. 42.
Langer Daumenstrecker. Tiedemann, § 274.

Prechtl, § 63.
Langer oder innerer Daumenstrecker. Heusinger, p. 190, No. 28

(Extensor pollicis).
Ohne Namen. Meckel, p. 349, No. 3.
Abductor 'pollicis brevis. Selenka, p. 138, No. 73.
Abductor ])ollicis. De Man, No. 37.
L'abducteiir direct de l' appendix. Court abducteur palmaire. Alix,

p. 418.

" Entspringt fleischig von der Yentralflache der Sehne des
M. extensor metacarpi radialis etwas proximal von dessen Insertion
an dem hervorstehenden Knorren des Metacarpale I. Der ziemlich
rundliche Muskel windet sich an der Yolarflache jenes Knorrens
vorbei und inserirt sich mit kurzer Sehne an dem radialen oder
vorderen innerer Yorsprunge des Pollex oder etwas distal da von "
(Bronn's Klassen, vi. Band, p. 289).

L 2

14S THE MYOLOGY OF THE RAYEN.

tomists for the foot, but it is incorrectly termed, as it
should be called the extensor proprius haJh.icis.

86. The fiexo)' hrevis i^oUicis arises from the shaft of
the mid-metacarpal bone of the carpo-metacarpus, just
below the anchylosed first or poUex metacarpal. Its
fibres converge as thev pass down behind the thumb
joint, to become converted into a delicate tendon which
is inserted into the distal apex of that phalanx.

Thus we see that the pollex of the Eaven is supplied
with two extensors and one fiexor. When the wing is
fuUv extended and the tendon of the extensor metaearpi
radialis longior is on the stretch, and at the same time
the tendinous slip from the extensor digit orunt com-
munis is pulling on this digit, it is held in full extension,
bY virtue of the several methods of attachment of the
muscles, and the wing feathers attached to it are in this
way so spread as to present the greatest amount of
superficial area to the atmosphere. When the wing is
closed, the more delicate flexor of the thumb draws that
joint down again towards the metacarpus, and is
sufliiciently powerful to retain it in that position.^

87. The jiexor minimi digiti^" instead of going to
the little finger as in a five-fingered hand (as in Homo),
it is inserted into the smallest finger-joint of the bird's
hand, which, as we know, corresponds with the medius.

1 \Ye find, upon referring to Selenka's excellent work in Bronn's
Tltier-Reichs (vi. Band, p. 138, No. 71), that he designates this muscle
as the Jiexor poUicis brevis, while Gadow describes it briefly as the

M. ADDUCTOR POLLICIS (No. 101) {loc. cit., pp. 291, 292).

- Here we have the muscle that Watson defined as the Jiexor
minimi digiti (p. 99), and which has been amply recognized by other
writers. Gadow made it his (No. 102) m. flexor digiti hi., and
remarked that " Bei Corvidee ist der Muskel nur schwach eut-
wickelt und, wenn iiberbaupt vorhanden, mehr auf den dorsal-
ulnaren Rand beschriiukt " {loc. cit., p. 293).

THE MUSCLES OF THE UPPER EXTREMITY. 149

It arises fleshy from the posterior aspect of the meclius
metacarpal close up to the os magnum, with a few fibres
extendiDg to the ulna (Fig. 44), and these two portions
converging, they soon send down the back of the carpo-
metacarpus, in a groove intended for it, a delicate tendon
which is inserted with the posterior margin of the base
of the medius phalanx.

88. The Jtexor minimi digiti hrevis'^ is in a very

^ Neither this muscle nor the next, my No. 89, is apparently
described by Professor Gadow, and, indeed, I fail to iind them
recognized by other writers on the subject, unless it be Selenka.
On the other hand, I find Gadow giving two muscles (herewith
reproduced below) that were not revealed to me during my dissec-
tions upon the Raven : —

" 96. M. ABDUCTOR INDICIS.

Vaddxicteur de la 2)remiere phalange. Cuvier.

Strecker des ersten Fingergliedes. Wiedemann, p. 94.

Strecker des ersten Gliedes des zweiten Fingers. Tiedemann ;

Heusinger, p. 193, No. 34.
Anzieher des zweiten Fingers, Meckel, p. 350, No. 8.
Anzieher des Zeigefingers [Adductor phalangis ^:»rma; indicis)

Schopss, p. 170, No. 45.
Niederzieher des grossen Fingers. Prechtl, § 61.
Court extenseur du medius. Milne- Edwards, pi. 10, fig. 1.
Vierter Interosseus. Selenka, p. 1 37, No. 70.
Interosseous IV. De Man, No. 36.
L'abducteur du deuxieme doigt. Gervais et Alix, p. 29.
L'abducteur palmaire du deuxieme doigt. Alix, p. 419.
Abductor digiti secundi. Watson, p. 100.
Abductor indicis. Carlsson, p. 28.

" Dieser Muskel kommt fleischig von der Radialflache des Meta-
carpale II. und zwar am gewuhnlichsten von dessen proximalem
Drittel dabei auf die Ventralseite iibergreifend. Bei besonderer
Entwicklung erstreckt sich der Ursprung auf die ersten drei Viertel
(Tauben, Kaubvogel) oder wohl auch auf nahezu die ganze Lange
jenes Knochens (Anser, Ibis, Psittacus), jedoch kann er, obgleich

150 THE MYOLOGY OF THE RAVEN.

nidimentaTy condition, tlioiigli we have no trouble in
making it out.

It arises from the lower and posterior end of the
medius metacarpal, and from the adjacent tendon of the
preceding muscle ; at least its meagre carneous portion
does, but its rather strong tendon, which has the same
osseous origin, is continued directly to its point of
insertion, the apex of the medius digit. So that in
function this muscle in the Eaven is rapidly coming to
play the part oi n posterior ligament to this joint, and
securely splices this now almost useless little finger to
the next digit by aid of the surrounding tissues, and
thus keeps it out of harm's way.

auf die proximale Halfte beschriinkt, doch i-echt stark sein z. B. bei
Bucorvus und Podargus

" Insertion. Die kurze starke Sehne verliiuft an der lunen-
Vorder Kaute des Metacarpale II. und inserirt sich am Innen-
Vorderhocker der Basis phal. I. dig. II."

In referring to Selenka's work in Bronn's Thier-Reicha (vi. Band,
p. 137), as noted in the above synonymy, I find that that anatomist
defines this muscle, his No. 70, as the abchicior diyiti ynimiai, and I
have every reason to believe that this muscle corresponds with my
No. 89, the abductor minimi digiti.

"97. M. FLEXOR POLLICIS.

Anzieher des Daumens {Adductor pollicis). Wiedemann, p. 94
Schopss, No. 43; Heusinger, p. 191, No. 31 ; Selenka, p. 138,
No. 74.

Anzieher oder Einwiirtszieher [Adductor ■pollicis). Tiedemann, § 282.

Ohne Namen. Meckel, p. 349, No. 4.

Einwartszieher des Daumens (pt.). Prechtl, § 65.

Lea courts Jlechisseurs du pouce. Alix, p. 418.

Flexor 2)ollicis brevis. Carlsson, p. 28." (Bronn's Klassen, «S:c., p.
288.)

Gadow also describes another muscle (No. 100 of his list), the
m. extensor pollicis brevis (lac. cit., p. 291), of which he says that
" So wird er bei den Corvidoi von Schops als nicht gefunden, von De
Man als zweifelhalf angegeben."

THE MUSCLES OF THE UPPEll EXTREMITY. 151

89. The abductor tninimi digiti is a small muscle,
almost reduced to a ligamentous baud, which arises by
a rather strong tendon and a few indistinct fibres from
the proximal extremity of the anterior aspect of this
medius digit, to pass downwards and become inserted
upon the posterior border of the proximal phalanx of
the index finger, at a point rather above the tip of the
medius digit. The action of this now highly rudi-
mentary muscle is very feeble, but, sucli as it is, it is
opposed to the more powerful flexors applied to the
back of this. joint.

90. The Jiexor riietacaiyi hrevis^ arises partly fleshy
and partly tendinous from the outer side of tlie distal
extremity of the ulna ; the carneous portion ceases at
a point about opposite the base of the thumb-joint.
From this point a delicate tendon passes obliquely down
the carpo-metacarpus to become inserted at the base
of the proximal phalanx of the index digit, on its ulnar
side, in front.

1 This muscle lias been defined as the tji. ulni-metacarjyalis dorsalis
by Gadow, who records the following synonymy for it {loc. cit., pp.
273, 274) :—

" 87. M. ULNI-METACARPALIS DORSALIS.

Le court Jlechisseur de I'os inetacarpe. Vicq d'Azyr, p. 577, No. 3.
Flexor metacarin h-evis. Tiedemann, § 273.

,, ,, „ Watson, p. 96.

„ ,, „ Heusinger, p. 190, No. 27.

Unterer Theil des iiusseren Ellenbogenmuskels. Meckel, p. 335,

No. 4.
Kurzer Beuger der Mittelhand. Schopss, p. 156, No. 36,

„ „ „ ,, Prechtl, § 56.

Flexor carpi radialis. Selenka, p. 134, No. 65.
C ourt Jlechisseur de la main. Milne-Edwards.
Le court adducteur de la main. Gervais et Alix, p. 28.
Court Jlechisseur du metacarpe. Alix, pi. II. fig. 2, No. 21.
Second ou court cubital p>osterieur, ou court adducteur de la main.
Alix, p. 140."

152

THE MYOLOGY OF THE RAVEN.

Fig. 42. — Anterior aspect of the hones oF the left hcand of a Raven, with thi;

pollex digit removed ; designed to show the origin and insertion of

some of the muscles or their tendons.
Fig. 43. — Uhiar aspect of the same bones; the axis of the exten^oi- proprius poIUcis

muscle is shown, and its origin indicated upon a piece of the extensor

mctacarpi rcflialis lonr/ior, given above.
Fig. 44. — Posterior aspect of the same bones. The several segments in all the

figures are slightly dislodged in order to show them to better advantage.

Siiecimens all life-size, and drawn by the author.

The action of this musch? is to flex the hand upon the
forearm, and at the same time rotate it slightly to the
inner side.

THE MUSCLES OF THE UPPER EXTREMITY. 153

91. The interosseous dorsal is is the outermost and
hisfher of the two delicate muscles that fill the long vacuity
existing between the index and medius metacarpal.
It is attached to the margins of the shafts of both these
bones, then sends obliquely downwaixls a slender tendon,
which lies fiat ao^ainst the radial side of the skeleton of
the hand, to be finally inserted at a point on the an-
terior aspect of the base of the distal phalanx of the
index dioit. The contraction of this muscle extends

o

the lowermost finger-joint of the hand in the Raven.

92. The interosseous pahnaris ^ lies in the same vacuity,

1 Gadow offers the following synonymy foi' the two interosseous
muscles I describe above, to wit : —

" 95. M. INTEROSSEUS DORSALIS.

L' interosseux anterieur. Vicq d'Azyr, p. 577.

Aeusserer Mittelhandmuskel (Interossetis externus.) Tiedemann,

§ 283 : Heusiuger, p. 197, No. 39.
Innerer 8trecker und Abzieher des zweiten Fingei's. Meckel, p.

350, No. 6.
Abzieher des Zeigefingers {Abductor ^j/i«/«?i^?s sccunclce indicis).

Schopss, p. 172, No. 46.
Oberer Mittelhandmuskel (Vorleger der Lenkfeder.) Prechtl, § 66.
Interosseux anterieur ou abducteur du medius. Milne- Edwards, pi.

9, fig. 3.
Interossevs. Selenka, p. 137, No. 69.
Jnterosseus II. ulnar is. Ue Man, No. 34.
Interosseux dorsal. Gervais et Alix, p. 30.
Vadducteur dorsal du deuxieme doigt. Alix, p. 419.
Interosseus dorsalis. Watson, p. 100 ; Carlsson, p. 25.

" 99. M. INTEROSSEUS PALMARIS.

U interosseux x>oslerieur. Vicq d'Azyr, p. 577.

Innerer Mittelmuskel {Interosseus internus). Tiedemann, § 284 ;

Heusinger, p. 197, No. 40.
Aeusserer Strecker und Abzieher des zweiten Fingers. Meckel, p.

350, No. 7.
Beuger des Zeigefingers. Schopss, p, 173, No. 47.

154 THE MYOLOGY OF THE RAVEN.

beneath the preceding muscle, while its fleshy portion
extends somewhat lower down in this intermetacarpal
fenestra to which it is attached than its companion.
Its tendon passes down in a manner similar to that
described for the interosseous dorsalis, but is carried
completely to the distal apex of the lowermost joint of
the index finger, being next to its posterior margin.
When this delicate little muscle contracts it flexes to a
marked degree the joint to which it is attached, and if
its contraction is carried to the full extent of which it is
capable, it will also flex both the digital joints of the
index finger, provided the opposition of the more power-
ful extensors is not applied at the time.

Unterer Mittelhandmuskel (Zuriickzieher der Lenkfeder.) Precbtl,

§67.
Interosseux posterieur ou court fleclnsseur du viedius. Milne-Edwax'ds,

pi. 9, fig. 3.
Interosseus. Selenka, p. 137, No. 69.
Interosseus I. radialis. De Man, No. 33.
Interosseux j^driiaire. Gervais et Alix.
Vadducteur du detixieme doigt. Alix, p. 419.
Interosseus palmaris. Watson, p. 101." {]i\-oi\D! a Klassen des Thier-

Reichs, vi. Band, p. 290.)

VIII. THE MUSCLES OF THE LOWEK
EXTREMITY.

To expose these, extend the median dorsal excision
through the integuments, to the very tip of the coccyx ;
join this by an extension of the median integumental
ventral incision. Next, join these two lines anteriorly
by a lateral division of the integuments, which above
passes immediately behind the point of the scapula.
Reflect the skin thus set free, and carefully remove it
as far as the ankle-joint. Here we meet with the
horny podotheca covering the tarso-metatarsus and toes.
This is best removed by lateral incisions and peeling
backwards and forwards both ways, being particularly
careful not to injure the tendons.

The following superficial muscles of the thigh are
now to be examined: —

93. The sartorius. 10]. The accessory semitendin-

94. The gluteus primus. osus.

95. The gluteus niedius. 102. The semimembranosus.

96. The gluteus minimus. 103. The femoro-caudal.

97. The extensor femoris (the 104. The obturator externus.

crurseus, the vastus ex- 105. The obturator internus.

ternus.) 106. The gemellus.

98. The vastus internus. 107. The abductor longus.

99. The biceps flexor cruris. 108. The addvictor magnus.
100. The semitendinosus. 109. The gastrocnemius.

156 THE MYOLOGY OF THE RAVEN.

110. The soleus. 118. The flexor perforatus annu-

111. The peroueus longus. laris primus pedis.

112. The tibialis anticus. 119. The flexor perforatus inedi us

113. The extensor longus digi- primus pedis.

torum. 120. The flexor perforatus medius

114. The extensor hall ucis bre vis. secundus pedis.

115. The tibialis posticus. 121. The flexor perforatus indici

116. The flexor perforatus indicis primus pedis.

secundus pedis. 122. The flexor perforans digi-

117. The flexor longus hallucis. torum profundus.

93. The sartorius ^ constitutes the most anterior of the
group of superficial muscles upon the outer aspect of
the thigh in the Raven, as it does in the vast majority
of birds.

It arises by a fleshy origin from the outer two-thirds
of the superior surface of the raised emargination of the
anterior border of the ilium, and by a few fibres and an
extension of its fascia, from the crest of the neural
spine of the fourth dorsal vertebra (Fig. 24). The muscle

^ Gadow has given us a very excellent account of this muscle for
birds, together with the subjoined synonymy, to wit : —

" 33. M. ILIO-TIBIALIS INTERNUS S. SARTORIUS.

M. quartus tibiam viovens. Aldrovandi.

M. irrimus femoris. Steno.

Le couturier. Vicq d'Azyr, p. 272, No. 1 ; Cuvier, p. 524 ; Gervais

et Alix, p. 30; Alix, p. 438.
Ausstreckender Schienbeinmuskel. Merrem, p. 159, No. 1.
Sartorius, Wiedemann, p. 94.

Tiedemann, § 294 ; Gurlt, p. 28.
„ Owen, Apteryx, p. 292.

„ Quennerstedt, p. 23.

„ Neander, p. 15 ; Selenka, p. 142, No. 82.

De Man, p. 125, 11,
„ Gadow, No. 13.

Watson, p. 109.
Rectus femoris. Ileid, p. 143." (Bronn's Klassen, vi. Band, p. 149.)

THE MUSCLES OF THE LOWER EXTEEMITY. 157

becomes much thicker and bulkier as it descends down-
wards and backwards towards the knee, its anterior
margin forming the free edge that bounds the group of
thigh-muscles in this reejion.

It is inserted, rather obliquely, by a semitendinous
fascia into the inner edge of the ligamentum patellae, and
by a somewhat stronger attachment to the inner and
contiguous border of the summit of the tibia for its
anterior half.

The sartorius is one of the extensors of the leg, it also
flexes and to some extent adducts the thio-h.

"We find the sartorius muscle (Fig. 62 his) power-
fully developed in Geococcyx, as are the majority of the
muscles of the thigh in that bird. It arises, semiten-
dinous, from the crest of the neural spine of the last
vertebra of the dorsal region of the spine, from the
summit of the anterior portion of the crista of the
sacrum, and from the adjacent surface of the superior
aspect of the ilium on the corresponding side. The
fibres, forming an oblong and rather thick muscle,
pass downwards and backwards to the region in front
of the knee. Here it becomes inserted by a special
slip of fascia that is thrown oft' and merges with
the general fascia surrounding the knee-joint ; and,
secondly, by a more carneous insertion into the inner
half of the superior rim of the cnemial crest of the
tibia and the continuous inner margin of the summit
of that bone.

" The sartorius in this bird bounds anteriorly the
superficial group of muscles of the thigh ; consequently
its anterior border is free. Its posterior border above
unites quite intimately with the overlapping gluteus
primus muscle ; while this border below is sharp and
free, although here, too, the gluteus also overlaps it.

158 THE MYOLOGY OF THE RAVEN.

and a delicate connective tissue binds them together."
(See 120 of Bihliogrwphy.)

94. Tlie gluteus primus is the most superficial of all
the muscles of the thigh, and but requires the removal
of the skin to bring it into full view. It overlaps the
preceding muscle in front and a number of the deeper
ones behind.

It arises by a thin fascia for nearly the entire length
of the supero-internal margin of the pre-acetabular portion
of the ilium, also from this bone above the anti tro-
chanter, and by a stronger origin from the entire length
of the postacetabular ridge. The anterior portion of the
muscle is thin and fascia-like, while behind it is better
developed. In front it forms a delicate covering to the
hinder moiety of the sartorius, and the body of the
gluteus medius ; behind, the biceps is the principal
muscle covered by it. From this long line of origin, the
fibres of this triangular muscle converge as they pass
downwards toward the patella, and just before reaching
that sesamoid they merge with the fibres of the extensor
femoris.

The combined muscles then form an extensive, thin
aponeurosis, which spreads over the anterior aspect of
the knee, and is inserted into the crest of the cnemial
border of the tibia, the well-developed patella being found
in this aponeurotic ligament.

In his earlier papers. Professor Garrod called this muscle
" tensor fasciw^' ; and Sir Richard Owen, speaking of it
in the Apteryx, says that " the most superficial of the
muscles on the outer side of the leg is that very broad one
which combines the functions of the tensor vagince and
rectus femoris, but which, in the opinion of Cuvier and
Meckel, is the homologue of the tensor vagincp. and gluteus
maximus (seu externusy (Anat. of Verts., vol. ii. p. 99).

THE MUSCLES OF THE LOWER EXTREMITY. 159

111 the Raven its anterior fasciculus can be separated
almost as a distinct muscle, and this portion has its distal
tendinous slip inserted more particularly over the anterior
surface of the patella.^

^ By the older anatomists this gluteus jmmus muscle was con-
sidered in its entirety in their descriptions, and it has, as will be
seen, so been considered here. More recent writers, however, and
very properly so perhaps, have divided this complex muscular mass
into three portions, of which the anterior portion is generally taken
to represent the recttis femoris ; the middle portion, the tensor fascice
latce ; and the hinder division a " m. glutoeus jiosteriorr

Gadow has admirably handled the subject, and the entire muscle
is claimed by him to be an Ulo-tihialis, and capable of division into
three parts. From this authority I here below obtain my synonymy,
and likewise republish largely from his accovint (see Bronn's Tliier-
Reichs, vi. Band, pp. 151-153) : —

" 34. M. ILIO-TIBIALIS.

" A. Der ganze Muskel-Complex.
M. primus tibiam moveris. Aldrovandi.
M. secundus femoris. Steno.
Latissimus /emoris. Wiedemann, p. 94.
Latissimus femoris = Tensor fasciae latoi. Tiedemann, § 295.
Tensor fasciae latce. d'Alton, p. 33.

Tensor fasciae et caput longum m. bicipitis femoris. Gurlt, p. 28.
Rectus femoris et tensor fasciae. Quennerstedt, p. 24.
„ „ ,, ,, Neander, p. 16.

" B. Die einzelnen Theile.
"I. M. ilio-tihialis anterior.
M. rectus femoris. Owen, Apteryx, p. 292.
Selenka, p. 14 2, No. 83.
DeMan, p. 124, No. 10.
., ,, Watson, p. 110.

M. ilio-tihialis externus. Gadow, No. 14.

" II. M. ilio-tibialis medius.

M. du fascia lata. Yicq d'Azyr, p. 272, No. 2 ; Cuvier, p. 523.
M. tensor fascicB lafca. Meckel, System, p. 360, No. 1 ; Gadov/,
No. 22.

160 THE MYOLOGY OF THE RAVEN.

95. The gluteus medius is a very difFercnt nuiscle
from the one just described, and in regard to it anatomists

Auswartswender und iiiisserer Beuger. Meckel, Archiv, p. 259,

No. 1.
Schenkelbindenspanner. Meckel, Archiv, p. 259, No. 1.
Tensor vagince. Owen, Apteryx, p. 292.
Selenka, p. 142, No. 84.
De Man, p. 124, No. 9.
I'tnseur du fascia lata. Gervais et Alix, p. 110.
Tensor fascioefemoris. Watson, p. 111.

" III. M. ilio tibialis jwsterior.

Grand fessier. Vicq d'Azyr, p. 272, No. 3 ; Cvivier, p. 523.
Grosser Gesassmuskel (hinterer Theil). Meckel, System, p. 361,

No. 1 ; Archiv, p. 259, No. 1.
Grand fessier. Gervais et Alix, p. 30 ; Alix, p. 430.
M. glutoius jwsterior. Gadow, No. 22.

" Die jiussere, oberflachliche Lage der Muskulatur des Oberschen-
kels wird von einer breiten, oft nur diinnen Mviskelmasse gebildet,
die drei verschiedene Muskeln repriisentirt. Da sie hinsichtlich
ihrer Ausbildung in Zahl und Ausdehnung die grosste Mannig-
faltigkeit zeigen, so wollen wir zuerst ihr typisches Verhalten
feststellen. Dieses finden wir bei den Hilhnern und Kranichien.
Die Muskeilage entspringt aponeurotisch vom ganzen dorso-lateralen
Rande des priiacetabularen und acetabularen Ilium, fern6r mehr
fleischig vom correspondirenden Kamme des postacetabularen Ilium,
dabei etwas auf das distale ende des Ischiwm iibergreifend, welolier
Theil dann vom M. caud. il.Jiexorius bedeckt wird. Die Muskelfasern
dieser breiten Schicht convergiren nach dem Kine zu, erreichen
dasselbe aber nicht, sondern heften sich mit einer starken Aponeurose
auf dem Endtheile des tiefer liegenden M.fem. tih. fest, verstiirken
mitliin dessen zur Patella tretende Sehne.

" Die hauptsachlichsten Verschiedenheiten sind folgende :
'' Erstens in Bezug auf die Insertion. Die ganze Muskelmasse ist
distal verkiirzt und inserirt bereits auf der Mitte des M. fevioro-
tibialis, ohne also das Kine zu erreichen {Ciconia, Ibis, Tauben,
Raubvogel, Papageien) ; oder sie reicht ziemlich bis zum Kine (die
meisten Sumpf-, Hiihner-, Schwimm-, und Singvogel). Der AI. ilio-

THE MUSCLES OF THE LOWEE EXTREMITY. 161

are; and generally have been, well agreed as to its
homology. It conies off from the entire supero-internal

tibialis anterior' ist dabei gevvohnlich der liingere. Bei Podiceps,
nicht jedoch bei Colyvibus, ist fast nur der mittlei-e, dem 31. ilio-tib.
■medius entsprechende Theil entwickelt ; dieser ist aber sehr breit
uad inserirt sich an der Patella, auch durch Verwachsung an der
Hinteraussenflache des M. gastrocnemius, dabei bis zur Mitte des
TJnterschenkels herabreichend. Bei den Sphenisciden inserirt sich
der mittlere und der sehr reducirte hintere Theil zvisammen mit dem
M. femoro-tibialis an der Aussenseite der Patella und am oberen
Eade der Crista tibite anterior externa.

" I. Haufiu; sind der Ilio-tib. und der Sartorius mehr oder weni^er
mit einander verwachsen, so besonders bei Pterocles, Columbm,
manchen Coccygomorphen {Rhamphastus, Podargus). ^eiSteganopoden,
Lariden, Aiken, und Chauna hingegen ist der Ilio-tib. vom Sartorius
durch eine grosse Liicke geti-ennt ; diese ist natiirlich hauptsachlich
dadurch hervorgebracht, dass entweder der Sartorius nur mit geringer
Ausdehnung vom Ilium entspringt, oder da.ss der Ilio tibialis anterior
sich auf den dem Acetabulum naheren Theil des Ilium beschriinkt.
Auch bei den Lamellirostren und den Sumpfvogeln bleiben beide
Muskeln meistens getrennt.

*' Am weitesten nach vom reicht der Il.-tib. bei Colymbus, denn er
entspringt ausser von dem ganzen Iliumknorren auch noch von den
Dorsalfortsatzen der letzten drei Riickenwirbel, auch inserirt er
nicht am M. femoro-tibialis, sondern direct an der Basis der Aussen-
kante der hohen Ci'ista tibio- patellar is. Hingegen fehlt der M. ilio-
tib. anterior ganz bei Phcenicopterus. Ganz frei und bandformig ist
er bei Bucorvus, entspringend vom Vorder- (proximal) Bande des
Ilium.

" Ganz allgemein ist der Ilio-tib. an seinem Hinterrande mit dem
Mitteltheile, dem Tensor fasciae, verwachsen, doch lasst er sich bei
manchen Sumpfvogeln, wie Grus, Crex, Numenius leicht davon
trennen.

" Bei Rhea ist er fast ganz vollstandig, und entspringt mit zwei
Kopfen. Der eine kommt mit platter Sehne zusammen mit dem
ventro-distalen Ursprunge des Sartorius (siehe dort) vom lateralen
Iliumrande, der andere plattsehnig von einem kleinen Theile vor
dem Processus acetabular is. Beide Kopfe sinde durch eine vom
dorsalen Iliumkamme kommende Aponeurose mit einander ver-
bunden.

M

162 THE MYOLOGY OF THE RAVEN.

margin of the pre-acetabular moiety of the ilium, and the
concave surface of the bone external to it. In this locality

" II. Der M. ilio tibialis medius oder M. tensor fasci(B. Diese
mittlere Masse ist weniger Variationen unterworfen. Sie beschriinken
sieh anf Reduction des mvisculosen Theiles, in dem Urspruug und
Insertion aponeurotisch werden. Dieser Theil muss aus folgendea
Griinden als eigener Muskel betrachtet werden.

" Obgleich der mittlere Theil der Innervation nocli zum M. il.-tib.
anterior gehort, ferner gewohnlicli mit demselben untrennbar
vereinigt ist, so ist er docli bei manchen, wie z. B. bei den Ratitea
vom Il.-tib. ant. getrennt, liingegen mit dem Glut. post, vereinigt,
Dass er in solchen Fallen wirklicli in dem als einheitlich erscheinen-
den Glut. post, enthalten ist, wird am sichersten durch die Innerva-
tion aus dem Cruralgebiet erwiesen.

" III. JUo-tibialis posterior oder Glutcaus posterior. Entspringt
fleiscliig vom laterodorsalen Kamm des postacetabulareu Ilium, und
zvvar vom grosseren Theile desselben bei den Katiten, Hiihuern,
Tauben, Pterodes, den meisten Sumpfvugeln, Ardea, Alectoriden,
Phcenicoj)terus, Lamellirostres, Colymbus, Podiceps, Cuculiden,
RJiamjjhastus, Pici und Capitoniden, und Passerinen. Bei Struthio
und Casuarius, ferner bei einigen Kasores und bei Cryptiirus reicht
er auf die Aussentiiiche des distalen Ischium, daselbst dann vom M-
caud.-il. flexorius bedeckt. Sehr kleiu, mit seinem Ursprunge auf
den Processus aceiabularis besclu'ankt ist er bei den S'ter/anopoden,
Spheniscidoi, Aiken, Moven, Sturmviigeln, Papageien, Raul)voge]n
(ausgenommen Cathartes), bei Upupa, bei den Goccygomorphen mit
Aiisnahme der Cuculiden, Ramphastido} und Coraciidoi. Bei den
meisten Coccygomorj)hen, fei-ner bei Giconia, Chauna, Sterna, den
Eulen, und manclien Papageien ist sein Vorhandeusein nur dadurcb
nachzuweisen, dass ein sehwacher Nerv aus dem Iscbiadicus sich
zu der Hinterfljiche der den Schenkel deckenden Muskelmasse
begiebt. Bei Buceros und Podargus endlich scheint ein Glutmus
jMsterior wirklich ganz zu fehlen."

For an instance of an author considering the entire gluteus p/rimus
to be the tensor fascia, we have but to quote the following paragraph
of Garrod's from his C ollected Scientific Memoirs, p. 189. lie says : —

" Tensor fascice. — This is the superficial muscle of the outside of
the thigh, covering the femur. It is flat and triangular in shape, and
arises as a membranous expansion which covers the gluteus ii., from

THE MUSCLES OF THE LOWER EXTREMITY. 163

it lies immediately beneath the attenuated, anterior
jDortion of the gluteus primus.

The fibres of the thick and fleshy muscle thus
formed fill in the concavity of the pre-acetabular division
of the ilium, converge and pass towards the outer aspect
of the proximal extremity of the femur, where they
become inserted obliquely upon the trochanter of that
bone, by a strong tendon, which rides over a bursa on its
anterior rim.

This muscle completely hides from view the gluteus
rtiinimus which is found beneath it.

96. Tlie gluteus minimus'^ is a far smaller muscle

the lower two-thirds of the posterior border of the iliac fossa in which
that muscle is situated, and from the fibrous septum which separates
that muscle from the gluteus iii. Further down it has origin also
from the whole length of the ridge which separates the postacetabular
area from the external lateral surface of the ischium, and which may
be termed the postacetabular ridge, as well as from the posterior border
of the ischium, as far forwards as its junction with the pubis, being
here slightly overlapped by the semitendinosus. The fibres converge
towards the knee ; and the deep portion of the muscle blends in its
course with the vastus externus, together with which it continues
forward to become part of the broad thin tendon which covers the
knee and is inserted in the front of the tibia-head, the patella being
situated in it, together with the long, slender, and flat tendon of the
amhiens muscle, which is situated below it, running obliquely from
inside and above, outwards and downwards. In many birds, as the
Falconidse and Psittaci, this muscle does not extend below the level
of the femur, but ends inferiorly by blending with the vastus
externus ; and consequently where such is the case, it evidently
cannot, as it does otherwise, cover any of the flexors of the le"". In
the Bucerotidse it is entirely absent. Whether this postacetabular
portion of the tensor fasciae is present or absent has some bearing on
classification, as in the different families it is a very constant
feature."

^ I add below the synonymy of the gluteal muscles from Gadow,
which will greatly assist the reader in comparing the muscles I
describe in the present work as the gluteus medius and y. mimirnus

M 2

164 THE MYOLOGY OF THE RAVEN.

than tlie gluteus inedius, and as I have just said lies
immediately beneath it. It makes a semitendinous

with the same muscles as they have been regarded and named by
other anatomists.

" 30. M. ILIO-FEMOKALIS EXTERNUS.

M. tertius femoris. Aldrovandi,

Le muscle pyra/ziiVZa/. Vicq d'Azyr, p. 273, No. 8 ; Cuvier, p. 503.

Pyramidenformiger Muskel. Merreni, p. 158, No. 5.

Pyramiden- oder birnformiger Muskel (J/, pi/rlformis). Tiedemann,

§ 288 , Gurlt, p. 27.
Oberer Zwillingsmuskel, oder eigentlicher Auswiirtszieher. Meckel,

p. 354, No. 4.
Auswiirtszieher des Oberschenkels. Meckel, Archiv, p. 262, No. 5.
Glutmus externus. Owen, Apteryx, p. 290.

„ „ Selenka, p. 139, No. 75.

De Man, p. 120, No. 1.
Glutmus [maxirnus). Quennerstedt, p. 13; Neander, p. 11.
Abducteur superieur de la cuisse. Milne-Edwards.
M. glutceus anterior. Gadow, No. 21.

" Dieser sehr kleine und flache Muskel ist von dreieckiger
Gestalt, und entspringt mit breiter, fleischiger Basis von der
Seitenflache des Ilium in Hohe des Acetabulum. Sein Ursprung
erstreckt sich stets bis auf die Linea dorsalis ilei. Nach vorn wird
er vorn M. ilio-trochant jmsterior begrenzt, den er theilweise bedeckt.
Nach hinten grenzt er an den M. ilio-fibularis. Er selbst wird vorn
M. ilio-tihialis bedeckt. Seine platte, ziemlich starke Sehne kreuzt
die des Trochanter externus oder etwas weiter distalwarts. Der
Muskel halt daher hauptsachlich das Femur am Becken fest und
zieht es etwas nach aussen."

•■' 29. Mm. ilio-trochanterici.

" I. M. ilio-trochantericus posterior.

M. primus femoris. Aldrovandi.
M. quintus femoris. Steno.
Moyenfessier. Vicq d'Azyr, p. 272, No. 4.

,, ,, Cuvier, p. 500.

,, ,, Gervais et Alix, p. 31.

AHx, p. 430.

THE MUSCLES OF THE LOWER EXTEEMITY. 165

attacliment to the anterior margin of tlie outer border of
the ilium, and has a somewhat more fleshy origin from

Glutceus magnus. Wiedemann, p. 95.
,, ,, Tiedemann, § 285.

Ghitceus riiaximus. Gurlt, p. 27.
Mittlerer Gesassmuskel, oder erster Heher des Oberschenkels.

Meckel, System, p. 352, No. 1 ; Archiv, p. 261, No. 2.
Glutceus medius. d'Alton, p. 32.

Owen, Apteryx, p. 290 ; CydopKdia, p. 295.
Selenka, p. 139, ISTo. 76.
De Man, p. 120, No. 2.
Quennerstedt, p. 14.
Neander, p. 10.
Watson, p. 103.
M. iliacus externus j)osterior. Gadow, No. 9.

" II. M. ilio-trochantericus anterior.

M. secundus femoris. Aldrovandi.

Liliaque anterieur. Vicq d'Azyr, p. 275, No. 5.

Iliacus minor ; kleiner Huftmuskel. Merrem, p. 159.

Iliacus anterior. Wiedemann, p. 95.

Glutceus medius. Tiedemann, § 286 ; Gurlt, p. 27.

Vorderer oder Kleiner Gesassmuskel (pt.). Meckel, System, p. 353,

No. 2.
Zweiter Heber des Oberschenkels. Meckel, Archiv, p. 261, No. 3.
Petit fessier. Cuvier, p. 503.

„ „ Gervais et Alix, p. 31.
Alix, p. 430.
Glutceus alter s. minor. d'Alton, p. 32.

„ ,, ,, (pt.). Owen, Cyclopcedia, p. 295.

Glutceus minimus. Owen, Apteryx, p. 291.

„ „ Selenka, p. 140, No. 77.

„ De Man, p. 120, No. 3.

„ ,, Watson, p. 103.

Glutceus 'minor. Quennerstedt, p. 12.

,, ,, Neander, p. 10.

M. iliacus externus anterior. Gadow, No. 11.

" III. M. ilio-trochantericus medius.
M. tertius femoris. Aldrovandi.

166 THE MYOLOGY OF THE RAVEN.

the siipero-external surface of the last rib, which is seen
to be immediately beneath it. From these two points
its fibres are directed backwards, downwards, and out-
wards, slightly converge, then become tendinous, and are
finally inserted into the outer aspect of the upper third
of the femur, below the trochanter, between the two
bellies of the extensor femoris.

Professor Owen, after completing his description of
the three gluteal muscles in the Apteryx, says further
that, " A muscle, which may be regarded either as a
distinct accessory to, or a strip of, the preceding one
[gliitceus minimus], arises immediately behind it from
half an inch of the outer and inferior part of the ilium ;
its fibres run nearly parallel with those of the glutcBUS
minimus, and terminate in a thin flat tendon, which
similarly bends round the outer part of the femur, to be
inserted into the outer and under part of the trochanter
immediately below the tendon of the glnUeus medius.
This muscle and the preceding portion, or glutceus
minimus, are described by Professor Mayer under the

Petit fessier. Vicq d'Azyi% p. 273, No. 6.

Glutceus minimus. Tiedemann, § 287.

"Vorderer oder Kleiner Gesassmuskel (pt.). Meckel, Archiv, p. 261,

No. 4.
Glutceus minor. Owen, Cydopcedia, p. 295.
Accessory to the glut, minim. Owen, Apteryx, p. 291.
GlutcBus quartus. Owen, Comp. Anat., ii. p. 100.
„ ,, Selenka, p. 140, No. 77.

De Man, p. 120, No. 3.
M. iliacus externus medius. Gadow, No. 10." (Bronn's Klassen, vi.
Bd. pp. 140, 141.)

JVote. — This synonymy must be taken with some degree of
caution, for I find it wrong in several instances ; as, for one example,
Owen does not call the muscle glutceus quarttis in vol. ii., p. 100,
of his Comp. Anat. of Verts., and Selenka is also misquoted for the
g. minimus (see Bronn's Klassen, vi. Bd. p. 140). — R. W. S.

THE MUSCLES OF THE LOWER EXTREMITY.

167

names of gluUeus quartus and glutceus quintus, in the
Cassowary ; one of them is absent in most birds " [Anat.
of Verts., vol. ii. pp. 100-101).

The two smaller gluteals pull the thigh-bone forwards
while at the same time they abduct it.

The group of gluteal muscles are also powerfully de-
veloped in our Geococcyx califoiiiianus. In it " the

^^•^Z

Fig. 45.— Aiiterioi' aspect of left femur of a Raven, designed to show the muscles

that are attached to it.
Fig. 46. — The same hone seen from behind ; a, femoral head of the flexor

ppA'forans digitonmi 2)rofundus.
Fig. 47. — The same bone viewed from its inner side. All life-size, by the author,

from his own dissections.

gluteus primus (Fig. 62 his) constitutes that great and
rather complex muscle which makes up the central fleshy
portion of the outer aspect of the thigh. It arises by a
strong fascia from the summit of the co- ossified neural
spines of the anterior sacral vertebrae, and by carneous
fibres from the outer rim and under surface of the

168 THE MYOLOGY OF THE RAVEN.

wliorl-like, overarching portion of the ilium behind ;
and finally from the contiguous portion of the pelvis
over the antitrochanter, between these anterior and
posterior origins. In front the muscle consists first of a
strong layer of semitendinous fascia, which closely over-
lies the gluteus medius muscle beneath it, and overlaps
the sartorius anteriorly. The posterior origin and mid-
division become rapidly carneous and more massive as
we proceed in the direction of the caudal extremity of
the body. So that, where we find it arising from be-
neath the overarching part of the ilium behind, the
muscle fills about one-fourth of the convexity there
formed, the semitendinosus filling the remainder of this
curious cavity. The fibres of the strong, semitendinous,
muscular sheet springing from these several origins, or
rather along this continuous line of origin, now pass,
converging as they do so, towards the anterior aspect
of the knee-joint. The semitendinous portion anteriorly
becomes fleshy as it arriv^es along the outer pelvic mar-
gin, with which it is quite intimately connected. The
hinder division of the muscle remains thick and car-
neous until it comes to the knee-joint. Here all the
fibres again become tendinous and fascia-like, and,
uniting with a similar structure contributed by the
extensor femoris lying beneath it, the combined sheath
thus formed surrounding the well-developed patella,
closely invests the front and sides of the knee-joint, and
is finally inserted all round the anterior and externo-
lateral borders of the summit of the tibia.

" The gluteus medius muscle (Fig. 63 his) is found to
be strong and tendinous. It, as in all of the birds that
I have examined, fills the concavity of the pre-acetabular
portion of the pelvis, and here in Geococcyx extends
laterally much beyond the bone, as this bird has a very

THE MUSCLES OF THE LOWER EXTREMITY. 169

narrow pelvis anteriorly, wliile it demands tlie use of a
powerful set of gluteal muscles.

" The gluteus medius arises by a strong, flat tendon
from the superior surface of the outer moiety of the
anterior iliac margin, by a dense fascia from the entire
line bounding the pre-acetabular concavity, and finally by
fleshy fibres from the upper side of the ilium itself.
The fibres of the roundish muscle thus formed converge
as they pass to the caput femoris, and, just before ar-
riving at the bone, they terminate in a dense flat ten-
don, which, passing over a bursa, is inserted at a point
on the antero- external asj)ect of the femoral trochanter.

" The gluteus minimus (Fig, 63 his) is a very much
smaller muscle than the gluteus medius, and is found
immediately beneath it to its outer side. In form it is
oblong, and fully three times as long as wide. It arises
from the outer superior surface of the fore-part of the
ilium, and passing obliquely downwards and backwards
as a flat narrow band of fibres, it becomes inserted by
semitendinous ones on the outer asj^ect of the upper
third of the femur, just below the trochanter. This
muscle may also ride over a small bursa, just before it
arrives at its insertion" (the present writer in Proc.
Zool. Soc. of London, 1886; see 120 of Bibliography
at end of this volume).

97. Tlie extensor femoris^ constitutes the great
extensor of the leg upon the thigh.

1 Bearing upon the nomenclatural history of the extensor femoris
and vasti muscles we have the following from Gadow's work in
Bronn's Klassen des Thier-Reichs (vi. Bd. pp. 154, 1-55) : —

" 35. M. FEMOKI-TIBIALIS.

" Der I. und II. Theil.
M. secundus tihiam movens. Aldrovandi.
M. sedecimiis femoris. Steno.

170 THE MYOLOGY OF THE RAVEN.

It is distinctly divided into two well-defined portions,
viz. the vastus externus and the crurceus. That part

Le 'muscle crural {Vaste externe et interne.) Vicq d'Azyr, p. 276,

No. 1.
Innerer grosser Muskel. Meirem, p. 159, No. 2.
Cruralis oder eigentlicher Schenkelmuskel. Wiedemann, p. 95.
M. cruralis cum vasto externo et inter7io. Tiedemann, § 297.
Unterschenkelstrecker. Meckel, System, p. 368, No. 7.
Tiefer Unterschenkelstrecker mit dem ausseren grossen Oberschen-
kelmuskel. Meckel, Archiv, p. 268, No. 3 u. 4 ; p. 269, No. 6 u. 7.
Le ti-iceps crural. Cuvier, p. 523.
Extensor cruris anterior. d'Alton, p. 34.
Rectus femoris et vastus externus. Gurlt, p. 26.
Crurceus et vastus externus. Owen, Cyclopoidia, p. 296 ; Selenka, p.

144, No. 89 ; De Man, p. 127, No. 15.
Crurceus. Owen, Apteryx, p. 293.
Cruralis et vastus externus. Quennerstedt, p. 28.

,, ,, ,, Neander, p. 18.

31. fem.oro-tihialis. Gadow, No. 17.
Extensor cruris. Watson, p. 115.

" Der III. Theil.
M. quintus tibiam movens. Aldrovandi.
M. sepdecimus /emoris. Steno.
Le droit interne (t). Vicq d'Azyr, p. 278, No. 4.
Hinterer grosser Muskel (?). Merrem, p. 159, No. 3.
Rectus femoris internus. Wiedemann, p. 98.
Tiedemann, § 298.
Gracilis, oder innerer gerader Schenkelmuskel. Meckel, System,

p. 367, No. 6 ; Archiv, p. 269, No. 5.
Vastus internus. Owen, Apteryx, p. 294.
De Man, p. 128, No. 16.
,, ,, Qviennerstedt, p. 30.

„ „ Neander, p. 19.

,, „ Alix, p. 436.

Crural interne. Gervais et Alix, p. 31. Alix.
M. rectus femoris internus. Gadow, No. 16.
Gracilis. Watson, p. 115."

Note. — I believe parts I. and II. here represent my crurceus and
vasttis externus, while part III. represents my vastus internus (which
see). No. 98

THE MUSCLES OF THE LOWER EXTREMITY. 171

which seems to be the homologue of the vastus exteimus
arises by a tendon on the outer aspect of the shaft of the
femur, at the base of the trochanter, and by fleshy fibres
adown the same side of that bone, nearly to the
condyle.

The crurceus has a bulk fully double the size of the
vastus externus ; it arises by a tendon from the anterior
and prominent rim of the trochanter above, and by
coarse, somewhat individualized bundles of muscular
fibres, down the antero-external aspect of the shaft
of the femur. These two muscles and the gluteus
2:)rimus merge below with each other and into that
tendinous fascia which spreads over the front of the
knee-joint to be inserted into the cnemial crest of
the tibia, the fascia of the muscles at the outer side
of the leg, and has in it (in its usual position when
present, as it is here in the Raven) the well-developed
patella.^

1 The gracilis muscle does not occur in the Raven, but as the
ambiens it received no little attention at the hands of Garrod and
Forbes, the former anatomist using it extensively in his classification
of birds.

Mr. Forbes described the ambiens in the following words. He
said : —

"This muscle, vinlike the others to be subsequently mentioned, lies
on the lower or inner surface of the thigh. As generally developed,
it is a more or less slender fusiform muscle, which, arising from the
prffipubic spine or process of the pelvis, close in front of the acetabu-
lum, runs along the inner side of the thigh superficially, and then,
running slightly outwards, runs, as a thin tendon, in the fibrous
tissues covering the knee-joint (in some cases perforating the
2)atella) to the outer side of the leg, and terminates there by joining
one of the tendons of the superficial flexor of the toe?, the fiexor
■perforatus digitorum. The course of this muscle will be made clear
by the accompanying representation of it, as seen in a Touraco
{Corythaix erythroloplui) . In one or two cases (e.g., QHdicneinius ,

172 THE MYOLOGY OF THE KAVEN.

In Geococcjjx californianus the amhiens is con-
spicuously developed.

It arises from the apex of the prominent prepubic
spine of the j)elvis, and the fibres passing directly down
to the inner side of the femur, and parallel with that
bone, form a strong fusiform muscle. As it approaches
the patella it terminates in a small flattened tendon,
which, piercing the fascial envelo2ie of the knee-joint
below the inferior apex of that sesamoid, passes round
the joint, to become finally lost to the outer side and
opposite the summit of the tibia, where some of its
tendinous fibres merge with the fibres of origin of the
Jlexor perforatus digitorum, or, at least, with one of its
divisions.

The amhiens is overlain by the sartorius muscle, and
in the figure is brought into view only through the
aid of a small dissecting-hook and chain, which pull it
forwards in order that it may be better seen (see Fig.
64 his).

98. Tlie vastus intenius is a very distinct and well-
defined muscle in the Kaven, as it is in a number of other
birds.

Stringops) it tends to become obsolete after reaching the knee,
becoming lost in the capsule of the knee-joint. In all Passerine
birds, and some others, it is always absent " (Coll. Scientijic Papers,
London, 1885, p. 195).

Several centuries ago Aldrovandi in his writings called the amhiens
muscle the vi. tertius tibice ; it was the m. quindecimus femoris oi
Steno, and the le crural grele of Vicq d'Azyr. Blasius, as well
as Merrem, termed it the " Langer Beinmuskcl," while it was the
gracilis of Wiedemann, Tiedemann, Owen, Gui-lt, Gegenbaur, De
Man, Selenka, Quennerstedt, Watson, and Neander. Cuvier and
Meckel had long French and German names for it respectively, and
it would seem that it received its present accepted name of the
amhiens from Sundervall in 1855.

THE MUSCLES OF THE LOWER EXTREMITY. 173

It arises from a point just below the head of the femur
on the postei'o-interual aspect of the shaft, and is attached
down the bone in a straight line, increasing in width as
it nears the knee, as low as the internal condyle. It then
clears the joint to become inserted as a rather broad
tendon along the thickened inner border of the summit of
the tibia. This muscle is pointed above, and its fibres

Fig. 48. — Thigli of Touraco {Corythaix erythrolojjha) viewed from the inner side,
to show the ambiens muscle, arising from the prtepubic spine of the
pelvis (p), and running along to blend with one of the tendons of origin
of the flexor perforatus digitorum (f.p.). F, femur ; Pi, patella ; i.e.,
inner condyle of femur ; t, tibia ; b, biceps (cut short) ; s, sartorius
(also cut) ; e, e, extensor femoris ; sm, semimembranosus ; add,
adductores. N.B.- — The surrounding parts have been .somewhat dis-
torted from their natural positions to show better the course of the
ambiens. (By the author, after Forbes.)

diverge as they descend, thus forming a subconical mus-
cular mass, which stands out prominently from the femoral
shaft. The lower and outer aspect of the muscle is ten-
dinous, which tendon in passing down merges with its
tendon of insertion into the tibial head.

The vastus internus also constitutes one of the
extensors of the leg upon the knee, and is a powerful
auxiliary to the extensor femoris.

174 THE MYOLOGY OF THE RAVEN.

99. Tlie hiceps flexor cruris ^ is a single-headed
muscle among birds, as it is here in the Raven. It
constitutes the principal one of those muscles holding
the more anterior position in the group at the hack of
the thigh. It arises beneath the gluteus j^i'inius by a
tendinous fascia from the postacetabular ridge, extend-
ing between the antitrochanter and the anterior point of
insertion of the semitendinosus. In form this muscle is
flat and triangular, the fibres converging as they descend
towards the knee.

All of this group of rear- thigh muscles lie in a plane
or planes roughly parallel to the median, longitudinal
plane of the body. After passing the knee-joint the

^ Extraordinary it surely is to find the number of names that this
muscle has received at the hands of anatomists, inasmuch as it is
easily distinguished, and possessed of peculiar characters. No two
writers out of a dozen have given it the same name, and Gadow
has given the subjoined synonymy for it (Bronn's T/der-Meichs, vi.
Bd. p. 168) :—

" 39. M. ILIO-FIBULARIS.

31. oc'dvus tihiam movens. Aldrovandi.
Biceps. Vicq d'Azyr, p. 277, No. 3.

,, Cuvier, p. 523 ; Q.uennerstedt, p. 25.

Neander, p. 16 ; Selenka, p. 143 ; De Man, 126, No. 12.
Zugespitzter Wadenbeinmuskel. Merrem, p. 159, No. 5.
Flexor cruris anterior. Wiedemann, p. 96.
Flexor cruris jmiinis anterior. Tiedemann, § 299.
Aeusserer oder Wadenbeinbeuger. Meckel, /System., p. 361, No. 2 ;

[and] Archiv, p. 271, No. 10.
Flexor cruris Jibularis. d'Alton, p. 34.
Caput breve hicvpitis femoris. Gurlt, p. 34.
Biceps Jlexor cruris. Owen.
Pulsator. Sundevall.
Biceps femoral. Gervais et Alix, p. 32.

Alix, p. 440.
Biceps cruris. Gar rod.
Biceps femoris. Watson, p. 112.
M. ilio-fibularis. Gadow, No. 27."

THE MUSCLES OF THE LOWER EXTREMITY. 175

muscular part of the biceps is superseded by a strong,
round tendon, wliich, passing down between the muscles
of the fleshiest part of the upper and outer side of the
leg, becomes attached to a tuberosity on the external
aspect of the shaft of the fibula, about two centimetres
below its head.

Thus this muscle becomes a powerful flexor of the leg
upon the thigh, but it has associated with it another
contrivance, so that when the leg is flexed the weight of
the posterior moiety of the body is in part transmitted
to the lower third of the femur.

This contrivance consists in a tendinous loop, the
longer and at the same time the inner end of which is
attached just above the outer condyle of the femur, while
the shorter end merges with the fascia of the supcro-
median aspect of the outer head of the gastrocnemius, and
the deeper muscles immediately beneath it. The tendon of
insertion of the biceps passes through the bight of this
exquisite little arrangement, and, in addition to the use
already assigned to it, as Owen says, it enables the
biceps to effect a more rapid and extensive inflection of
the leg than it otherwise could have produced by the
simple contraction of its fibres.

Coming to consider these muscles (the biceps flexor
cruris, vastus internus, and the extensor femoris), in such
a form as Geococcyx calif ornianus, I have elsewhere re-
marked that " the hiceps flexor cruris (Fig. 63 his) arises
by carneous fibres upon quite an extensive portion of the
under surface of the over-curled part of the ilium behind
the acetabulum, and by a long tendinous slip w^hich comes
off from the free anterior margin of this part of the ilium.
The fibres converge as they pass downwards, and unite to
form a somewhat flattened muscle. Opposite the head
of the tibia, the biceps terminates in a round tendon, of

176 THE MYOLOGY OF THE RAVEN.

cord -like dimensions, which passes through a special loop
to make its way between some of the muscles at the
hack of the leg, to become inserted on the tubercle in-
tended for it on the outer side of the superior moiety of
the shaft of the fibula. The loop of the biceps is flat
and fashioned like a delicate tendinous ribbon. Its
upper end arises from the side of the shaft of the femur
above the external condyle, while the lower end comes
off from this protuberance just below the insertion of
the outer slip of the external head of the gastrocnemius
muscle, A branch of the sciatic nerve also passes
through this loop in company with the tendon of the
hice2:)s.

"The extensor femo)-is in readily divisible at its lower
half into two parts, the bulkier anterior one representing
the crurcBUs (Fig. 63 his), and the posterior division the
vastus externus.

" As a whole, this powerful extensor of the leg upon
the thigh arises from the antero-external aspect of nearly
the entire length of the shaft of the femur, and from a
portion of the trochanter at its summit. At about its
lower fourth it terminates in a broad tendinous expan-
sion, which, as has already been described, is amply re-
enforced by the tendon of the gluteus 2)i'iuius, which is
situated on the upper side ; the combined tendon thus
formed surrounds closely the anterior aspect of the knee-
joint, and is finally inserted into the proximal end of
the tibia, upon its front and external margins.

" The patella is found encased in front in this great
tendinous sheath of the knee-joint, and below the ajDex
of this sesamoid we find the enveloped track of the
tendon of the ambiens muscles, as it passes round in
front of the fcmoro- tibial articulation. The coml)ined
tendon of the extensor femoris is finally inserted into

THE MUSCLES OF THE LOWER EXTREMITY. 177

tlie cnemial crest of the tibial and the lateral boundaries
of the summit of that bone. Some of the superficial
muscles on the outer side of the leg are so extended as
to take a certain amount of their oriorin from this great
tendinous expansion.

" In (Fig. 63 his) I have very thoroughly divided these
two subdivisions of the extensor femoris, in order to
show their relative size, as well as their relation to each
other and the surrounding structures."

100. The semitendinosus ^ is abroad and flat muscle,
which, aided by the semimembranosus beneath it, forms

^ For interesting accounts of this muscle as it is found in others
of the class, see Selenka's myology of Aves (Bronn's Thier-Reichs,
vi. Bd. p. 143), and Gadow's excellent chapter on the same sub-
ject (loc. cit., pp. 162, 164). The latter writer has called it the
Caud.-ilio-flexorius, and both Selenka and Gadow have treated the
"Jf. accessorius se7nitendinosi " under the same muscle; the last-
named is ISTo. 101 of the present work. As heretofore, I republish
the synonymy of the semitendinosus from Gadow : —

" 37. M. CAUD-ILIO-FLEXORIUS.

M. sextus tihiam rnovens. Aldrovandi.

M. tertius femoris. Steno.

Le muscle qui tient la ^;?ace du demi-memhraneux ou du demi-

nerveux. Yicq dAzyr, p. 277, No. 2.
Hinterer Anzieher des Beines. Merrem, p 159, No. 4.
Flexor cruris posterior. Wiedemann, p. 96.
Flexor cruris tertius s. p)osterior. Tiedemann, § 301.
Le muscle demi-iierveux. Cuvier, p. 524.
Schienbeinbeuger. Meckel, System, p. 362, No. 3 ; Archiv, p. 269,

No. 8.
Seinitendinosus. Owen.

„ Selenka, p. 143.

„ De Man, p. 126, No. 13 ; Quennerstedt, p. 26.

„ Garrod ; Watson, p. 113.

Demi-tendineux. Gervais et Alix, p. 32.

„ Alix, p. 441.

M. caudi-ilio-Jlexorius. Gadow, No. 25."

N

-^jC^

flexor l^ri^tiff IvnllxiciS.

Pig. 49. — Skeleton of the left posterior extremity of an American Kavcn, seen
from the outer side. Drawn by the author from his own dissections,

THE MUSCLES OF THE LOWER EXTREMITY. 179

and designed to show the origin and insertion of a number of the
muscles of the hind limb. f.jJ.i.p.p., flexor pcrforatus indicis lyrimus
pedis.
Fig. 50. — Sketch of anterior asjiect of the head of the tibia of the same specimen,
enlarged ; showing insertions of the muscles there found. 1, the
extensor femoris ; 2, peroneus longus ; 3, tibialis anlicus (inner head) ;
4, extensor longus digitorum ; 5, separate slip from extensor longus
digitoi'um.

behind tlie muscular contour of the tliigli. It arises
from the hinder third of ^the postacetabuhir ridge, and
by a thin, though strong, fascia from the surface of the
caudal muscles beneath it. Its fibres pass obliquely
downwards and forwards to become inserted into the
tendinous raphe which is found along the posterior
truncate margin of the accessory semitendinosus, the
lower point of which raphe merges with the median
fascia of the inner head of the gastrocnemius muscle.

So far as I can ascertain from anatomical works at
present available, the origin of the semitendinosus in
the Raven seems to be, at least, quite an uncommon one.
Owen, speaking of it for the Apteryx says, it " arises
from the posterior and outer part of the sacrum and the
aponeurosis connecting it with the ischium;" while Forbes,
describing it more generally for the class, says that " it
arises from the transverse process of the first free coc-
cygeal vertebra, and from the fibrous membrane between
this and the ilium." Indeed, we have yet much to
investigate in the myology of this group of vertebrates !

101. The accessory seinitendinosus is a fiat oblong
muscle that arises from an oblique line just above the
condyles, on the posterior aspect of the shaft of the
femur (Fig. 46). Its fibres, passing directly upwards
and backwards, attach themselves to the tendiuous raphe
common to this muscle and the semitend/inosus, already
described in the preceding paragraph.

In the Raven the accessory semitendinosus has a

N 2

180 THE MYOLOGY OF THE KAVEN.

leno:th of sometliino; over two centimetres, with a corre-
spondiiig width of about one centimetre, while in thickness
it does not exceed tv/enty-five millimetres. The raphe
is continued on, as a slender tendon, to the inner side of
the shaft of the tibia, where it becomes inserted.

102. TJie seniimemhranosus^ (Figs- 24 and 50) also
contributes to the contour of the posterior line of the
fleshy part of the thigh in the Eaven. It is a long,
narrow, somewhat ribbon-like muscle, that arises from the
outer surface of the ischium of the pelvis, from the lower
margin of its notch on the posterior pelvic border and
extending on a curved line on the surface beyond,

^ As is well known, this is another muscle that Garrod describes
in his chapter on the value of muscles in classification, although this
one was hut little used owing to the fact that it is quite constant
in the class.

Gadow, who well describes it, gives us the following synonymy
{Joe. cit., p. 16G) : —

"38. M. ISCHIO-FLEXORIUS.

M. Septimus tihiam onovens. Aldrovandi.

M, quatuordeciri/MS femoris. 8teno.

Demi-nerveux {X). Vicq d'Azyr, p. 277, No. 3 (pt.).

Biceps (pt.). Yicq d'Azyr, 1774, p. 507, ISTo. 3. *

Kleiner Lendenmuskel ('?). Mei-rem, p. 158, No. 4.

Flexor cruris tertius. Wiedemann, p. 97.

Flexor cruris quartus. Tiedemann, § 302.

M. demi-mevibraneux. Cuvier, p. 523.

Ohne Namen. Meckel, System, p. 364, No. 4 ; Archiv, p. 270, No. 9.

Flexor cruris tibialis. d'Alton, p. 34.

Semimembranosus. Gurlt, p. 29 ; Selenka, p. 144.

„ „ (pt.) Quennerstedt, p. 27.

„ ,, (pt.) Neander, p. 17.

„ „ De Man, p. 127, No. 14; Garrod; Watson,

p. 113.
Le droit tnterne. Gervais et Alix, p. 32.

„ ,, Alix, p. 442.

M. ischio-Jlexorius. Gadow, No. 26."

THE MUSCLES OF THE LOWER EXTREMITY. 181

as shown in Ficr. 24. From this orimn this straight
muscle passes directly downwards and forwards, and
when it arrives at the muscles of the leg, it becomes
converted into a delicate, thin tendon which, passing
between the muscles of these parts, reaches the inner side
of the shaft of the tibia about one and a half centimetres
below its head, or rather bounding line of its summit,
where it is inserted on a line parallel with the long axis
of the tibial shaft (Fig. 50).

The semimemhranosus is a direct flexor of the lesr
upon the thigh ; it lies nearly in the same plane with the
sernitendmosus, the contiguous borders being joined by
the surrounding connective tissue.

Mr. Garrod says of these last three muscles that
" some birds, as in the Eagles and Owls, have no semi-
tendinosus at all ; some, as the Anserine birds and
Penguins, have no accessory semitendinosus, in which
case all the fibres go straight to the tibia-head ; whilst
in most the above-described condition maintains ; " and
of the semimembranosus he remarks that " this muscle
is very constant in birds ; in the Grebes it is extremely
thin, and may sometimes be absent, as stated by Sunde-
vall ; but I have seen it in some fresh specimens of
Podiceps minor, though but very slightly developed "
{^Coll. Scientific Memoirs, London, 1881, p. 190).

And of the same three, as they occur in Geococcyx,
I said in the P.Z.S. of 1886, that the semitendinosus
(Fig. 64 his) is a marvellously well-developed muscle in
this form, as is also its accessory head. Its origin fills
about three-fuurths of the nether cavity formed by the
posterior overarching portion of the ilium, under which
it arises.

Posteriorly, the fibres forming its free margin are so
arranged as to create a rounded border ; the lower end

182 THE MYOLOGY OF THE RAVEX.

of its arc terminating about opposite the post-pubis of
the pelvis. From this origin the fibres of the semiten-
dinosus pass downwards and forwards as a great, though
somewliat compressed muscle. When within rather
more than a (-enti metre's length of the shaft of the
feiDur, they terminate in an obUque tendinous raphe,
which latter forms the boundinor-line between this
muscle and the next.

The accessor// seinitenduiosus (Fig. 64 his) is composed
of coarser fibres than the muscle just described. It
springs from a longitudinal line occupying the distal
half of the shaft of the femur, and from the upper
surface of the hinder aspect of the external condyle of
that bone. The fibres pass backwards and a little up-
wards to become inserted into the tendinous raphe just
alluded to.

The lower extremity of this tendinous raphe termi-
nates, in Geococcyx, in a thin, flat, and delicate tendon,
which continues downwards and forwards to the inner
surface of the head of the tibia, where it becomes in-
serted, the point of insertion being found above that of
the semimemhixtnosus muscle, the insertional tendon of
which overlaps it.

The semimembranosus (Fig. 64 his) in Geococcyx,
though thoroughly developed, is rather a slender and
thin muscle, markedly so when we compare it with
the massive semitendiiiosus which overlies it.

It arises from the outer surface of the ischium, for its
posterior two-thirds, on a line situated a few millimetres
above the low^er free edge of that element of the pelvis.
The fibres gradually converge as they pass downwards
and forwards, to terminate in a very delicate and thin
ribbon-like tendon, wdiich, passing between the broad
tibial head of the gastrocnemius and the proximal

THE MUSCLES OF THE LOWER EXTREMITY. 183

extremity of the shaft of the tibia, becomes finally
tJiereuj^on inserted on its internal surface. The hinder
maroin of the semime^nhrauosus is free, while its border
anteriorly is juxtaposed to the posterior edge of the
adductors.

103. The feinoro-caudal ^ is another one of that
group of five or six muscles made historical through the

1 Gadow has carefully gotten together quite a complete synonymy
of this muscle for us, which shows, in the most interesting way
possible, the various christenings it has been at various times sub-
jected to. I reproduce this synonymy here, it being fi'om the work
so frequently quoted throughout the present treatise : —

" 36. M. CAUD-ILIO-FEMOKALIS.

M. sextiis femoris. Steno.

Le deuxieme ahducteur de la cidsse ; M. cruro-coccygien. Vicq

d'Azyr, p. 278, No. 3.
Schwanzhiiftmuskel. Merrem, p. 158, No. 2.
M. cruro-coccygeus + Adductor 2^i^nus femoris. Wiedemann, p. 96

u. 98.
Adductor jn'imus femoris + Cruro-coccygeus. Tiedemann, § 225 u. §

290.
Birnmuskel. Meckel, System, pp. 355-357, No. 5.
Biroenfbrmiger Muskel. Meckel, Archiv, p. 263, No. 8.
Le femoro-caudien du cruro-coccygien. Cuvier, p. 288, No. 8.
Geinellus superior + inferior. Gurlt, p. 20 u. 27.
Adductor longus fe-nioris. Owen, Apteryx, p. 291.
Abaisseur superieur de la cuisse. Milne- Edwards.
Triceps adductor femoris (third head). Reid, p. 143.
Femoro-caudalis + Caput pelvinum m.fe^noro-caudalis. Sundevall.
Adductor longus. Selenka, p. 141, No. 81.
De Man, p. 123, No. 8.
Femoro-caudal + Accessoi^y-femoro-caudal. Gari'od.
Femoro-coccygien. Gervais et Alix, p. 32.

,, ,, Alix, p. 433.

M. caudi-ischio-ilio-femoralis. Gadow, No. 24.
Adductor longus femoris + Cruro-coccygeus. Watson, p. 105.
Hierzu Kommen noch die kleinen Hiilfsmuskeln, in der Literatur

meistens ohne Namen.

184 THE MYOLOGY OF THE RAVEN.

numerous dissections of tlie tliighs of birds by the genius
of Garrod.

In the Raven, as in so many of the cLass, it is, next to
the biceps criiris, one of the most interesting muscles
that our scalpel reveals for us.

In form it is a long narrow spindle, flattened from
side to side. Its caudal extremity is drawn out into a
delicate tendon, which, arising from the base of the
pygostyle of the tail-skeleton, passes between the lateral
caudal muscles towards its insertion.

Its anterior extremity is also tendinous, but thin and
flattened in the same plane with the muscle. This is
inserted upon the outer aspect of the shaft of the femur,
below the trochanter, at about the junction of the upper
with the middle third of the bone. With the limb fixed,
and the muscle acting from its femoral end, it would
assist the lateral caudal muscles in drawing the coccyx,
and with it the tail, to its own side ; but with the coccyx
fixed, it would, in contracting from that end, tend to
pull the femur backwards and slightly rotate it out-
wards.

In describing the origin and insertion of this muscle
for the class generally, Garrod says that " it arises from
the (anterior) transverse processes of the two last coc-
cygeal vertebrse, and is inserted into the linea aspera
of the femur, at about one-third its length from the
trochanter." The fact that it arises from the base of the
pygostyle (Fig. 24) in our present subject forms quite
an interesting exception to the general' rule as laid down
by the talented anatomist just quoted.

Ohne Namen. Meckel, /System, p. 356, No. 5, unci p. 357, No. 7 ;

Archiv, p. 265, letzter Absatz vor No. 10, und p. 262, No. 6.
M. quadratus femoris ; Adductor hrevis. Owen, Apteryx, p. 292,

291." (Bronn's Klassen des Thier-Reichs, vi. Band. p. 158.)

THE MUSCLES OF THE LOWER EXTREMITY. 185

It may be as well to observe that the point of insertion
of tliis muscle upon the shaft of the femur is some httle
distance below that of the gluteus minimus ; the vastus
extenius of the extensor femoris passes up between
these two muscles, and the femoro-eaudal itself overlies
the adductor muscles which pass down between it and
the pelvis ; so that its insertion may also be said to be
on a narrow vertical line between the insertions of the
vastus externus and the adductors. ■"■

In my memoir upon Geococcyx I said that in that
genus " TliQ femoro-eaudal muscle and the accessory
femoro-eaudal are both present and fully developed.

" T\iQ femoro-eaudal (Fig. 64 his) arises, tendinous,
from the lower posterior border of the pygostyle. It
soon becomes fleshy, and as a narrow, muscular ribbon
passes through the tissues overlying the lateral group of
caudal muscles proper. Opposite the posterior border
of the pelvis it expands to form a prettily-shaped and
compressed spindle, closely covering the ohturator ex-
ternus muscle and the side of that bone. As it nears
the femur it again contracts, receives the fibres of its
accessory head, and is finally inserted upon the femoral
shaft, at the posterior aspect of its proximal third.

^ Many birds have an accessory femoro-eaudal ; it is absent in the
Raven. It is described by Garrod as " an accessory head, arising
from the upper three-fourths of the postacetabular ridge, and from
the ridge which forms the lower margin of the origin of the ob-
turator externus, joins the tendon of insertion of this muscle, and
is also partly inserted into the linea aspera, between it and the
head of the femur. It is thin, muscular, and broad, covering the
obturator externus superficially, and is partially intersected by a
fibrous sheet where it crosses its anterior border. The sciatic artery
and nerve cross it superficially ; and the nerve to the semimem-
branosus is deep of it, whilst that to the semitendinosus is superficial
in some cases ; the biceps completely covers it " {Coll. Scientifio
Memoirs, p. 191).

186 THE MYOLOGY OF THE RAVEX.

" The accessory femoro-caudal (Fig. 64 his) arises be-
neath the overarching part of the postacetabular portion
of the ilium, just behind the acetabulum and beyond.
Its fibres pass obliquely downwards and forwards to join
with those of the femoro-caudal, and to become inserted
with them into the upper part of the femur as already
described."

104. The ohtuvator exterims^ is a thick fleshy muscle
that arises from the posterior half of the periphery of
the ischiatic foramen and the concavity found on the
external surface of the lateral aspect of the pelvis be-
hind it ; its boundary above being the postacetabular
ridge, while its fascia is nearly carried to the posterior
pelvic margin behind (Fig. 24). From this origin it
passes directly toward the femur, its fibres converging

^ The subjoined synonymy is from Gadow, loc. cit., p. 170: —

" 40. M. ISCHIO-FEMORALIS.

21. quartus femoris . Aldrovandi.

M. duodecimus femoris. Steao.

Le vmscle qtd tient la place du quarre. Vicq d'Azyr, p. 273, No. 10.

Rotator fevtoris. Wiedemann, p. 97.

Obturator exteriius. Tiedemann, § 289.

Watson, p. 107.
Zweiter Niederzieher, oder viereckiger Schenkelmuskel, oder

ausserer Hiiftbeiuloclimuskel. Meckel, Sijstem, p. 357, No. 6 ;

Archiv, p. 265, No. 10.
Le carre de la cuisse. Cuvier, p. 503 ; Milne-Edwards.
Gemellus superior, d' Alton, p. 32.
Glutoius maximus. Coues, p. 168.
Quadratus femoris. Gurlt, p. 27.

„ „ Selenka, p. 140, No. 79.

De Man, p. 122, No. 6.
Pyramidalis. Owen, Apteryx, p. 291 ; Kuhl, p. 79.
Pyriformis. Quennerotedt, p. 19.

,, Neander, p. 13.

Le carre. Gervais et Alix, p. 32 ; Alix, p. 432.
M. ischiofeinoralis. Gadow, No. 23."

THE MUSCLES OF THE LOWER EXTREMITY.

187

SCwtbTWLS

■ J't 'u

Fi^.Sp.

Fis-^3'

Fig. 51. — Left tibia and fibula of a Raven viewed from the inner side.

Fig. 52. — The same bones seen from in front.

Fig. 53.— The same from a posterior aspect. Designed to show in each case the
origin and insertion of their muscles. In each, the patella is placed
in position above the tibia. All the drawings are life-size, from the
author's own dissections.

to form a broad, flat tendon, which is inserted into
the shaft of that bone just below its trochanter. This
insertion is almost directly opposite that of the gluteus

188 THE MYOLOGY OF THE KAVEN.

minimus, and tlie vastus externus muscle of the exten-
sor femoris is inserted on a longitudinal line between
them (Fig. 49). In the absence of the accessory femoro-
caudal as we find it here, the sciatic nerve and artery
passes over and external to the tendon of the obturator
externus, its track being impressed upon it in spirit
sj)ecimens.

In the course of my remarks on the value of certain
of these muscles in the classification of birds, published
in another connection (see 124 of Bihliography) I said
that, " There are Jive muscles in the thigh which have
proved to be more or less useful in the classification of
birds. These muscles are the following, and four of
them I have designated by the letters which were used
by Garrod in his myological formulae.

6. The ambiens,

7. The femoro-caudal A

8. The accessory femoro-caudal B

9. The scmitoidmosus X

10. The accessory seiyiitendinosus Y

" AVe know of no bird in which all five of these
muscles are absent, or' even of one which lacks the last
four in the list.

"According to Garrod, 'when these four muscles are
present in a bird, the formula AB. XY expresses the
fact ; when any one is absent, that such is the case is
indicated by the omission of the letter representing it.
Thus the formula A. XY indicates that the accessory
femoro-caudal muscle only is absent ; AB. X that the
accessory semitendinosus is missing; A.X that the
femoro-caudal and semitendinosus only are to be found ;
and A that the femoro-caudal alone is present.'

THE MUSCLES OF THE LOWER EXTREMITY 189

" This eminent anatomist applied these myological for-
mulae to a classification of the entire group of existing
birds, and fully discussed the matter in his work in the
most masterly manner in so doing, but it will be im-
possible to enter upon any such field here. In my own
opinion, however, I am inclined to believe that Garrod's
classification stands in need of a very thorough overhaul-
ing in many of its aspects ; by this I mean that in a vast
number of cases we are not in possession of the requisite
knowledo;e of the entire structure of certain forms as to
warrant one retaining them where Garrod has placed
them. In other words, these myological formulae, as
time goes by, and our knowledge of avian morphology
widens, will surely prove very useful in taxonomy, hut
they can only be employed with safety when taken, as
one set of characters, in connection with all the others
that the organization of any particular bird-form pre-
sents us with, and by no means are we to rely upon
them alone, or even when a few other sets of structural
characters seem to indicate a bird's affinity.

" To illustrate my point, let us turn for a moment to
the Swifts and Humming-birds ; here we have two
groups which for years past have been associated to-
gether as allied forms by systematists, and Garrod, too,
seemed to believe in their affinity. Why ? Because
the formula for the thio;h-muscles in each case was
found to be A ; the sternum had in each case an un-
notched posterior border ; and neither Swifts nor Hum-
mers possess intestinal caeca. Yes, this all may be so,
but all the rest of the organization of these birds is as
widely different as one can well imagine, and conse-
quently they belong to very different orders of birds.
This latter statement gains weight when we come to
think that aside from the formula for the thigh-muscles

190 THE MYOLOGY OF THE RAVEN.

being the same in C//pseli and Troclnh, their pelvic
limbs otherwise are by no means alike in other par-
ticulars ; and the sternum is, too, of a very different
pattern in each case, altliough, as I say, each possesses
an entire posterior xiphoidal margin.

" As in the case with the other muscles described in
the foregoing paragraphs, ornithotomists have a fine
field open before them Iq dissecting out this group of
thigh-muscles in our United States birds ; making full
notes upon their researches, and comparing carefully
with the W'ork already accomplished by the indefatigable
Garrod. In doing this, not merely the absence or pre-
sence of the five muscles last described should be noted,
Ijut, if possible, full notes made as to their exact origins
and insertions, their relative size as compared with
other allied Ijirds, and in short their morphology in its
details,"

105. Tlic ohturator internui^ ^ in the Eaven, as in most
birds, is a l)ipenniform muscle which arises from the
ventral surface of the ischium, as far Ijack as to include

1 Cuvier, even so far back as his time, considered this muscle to
be the " obtvratevr interne," and with but one or two exceptions, it
lias universally been so regarded since the earlier works of Owen.

Gadow gives the following synonymy for it, it being his 3f.
ohturator (loc. cit., 171) : —

'* 41. M. OBTUKATOR.

M. qiiintus fern oris. Aldrovandi.
M. decimus J'emoris. Steno.
L'iliaque interne. Vicq d'Azyr, p. 275.
Iliacus internus. Wiedemann, p. 98.
Tiedemann, § 29;3.
Dritter Anzieher, Einwartszieher oder Kammmuskel. Meckel,

System, p. 359, No. 10 ; ArcMv, p. 265, No. 11 u. 12.
Ohturator interne. Cuvier, p. 503.
Ohturatorius. d' Alton, p. 33.

THE MUSCLES OF THE LOWER EXTREMITY. 191

its hinder margin ; from the inner line of the corresponding
postpubis ; and from the membrane filling in the ob-
turator space between these two elements. It is of a
subtriangular form, flat, and possesses a central tendon
to which its fibres converge in an oblique anterior
direction. AVheii this tendon reaches the obturator
foramen it becomes dense, subcylindricnl, and strong,
and passing through this vacuity it is inserted, in
common wdtli the gemellus, into the outer aspect of
the trochanter of the femur, opposite the insertion of
the gluteus 7nedius. Outside the pelvis the tendon of
this muscle lies upon the gemellus and is external to it.

The late Professor Garrod made a point of calling
attention to the fact as to wdiether the area covered by
the origin of the obturator intenius was of an oval or
a triangular outline, and entered the information in his
valuable tables of the anatomical characters of birds.
It seems to me, however, that such a character as this
must bear with it but very little weight, as the form
of the muscle, so far as my observations go, varies
directly with the form of the pelvis, and particularly
with the form of the osseous elements to which it is
attached.

About three years ago, I published an article entitled
'■ A Keview of the Muscles used in the Classification of
Birds" (see 124 of the Bihliography at the end of this

Obturator internus. Owen, A2)teryx, p. 292 ; Reid, p. 143 ; Gurlt,
p. 28.

„ „ Garrod.

„ „ Quennerstedt, p. 14. '

,, „ Neander, p. 12.

„ „ Watson, p. 108.

Ahducteur interne de la cuisse. Milne-Edwards.
L' ohturateur externe. Gervais et Alix, p. 31 ; Alix, p. 434.
]\[. obturator. Gadow, No. 19."

192

THE MYOLOGY OF THE RAVEN.

volume), in which I made certain remarks upon " the
area of the origin of the obturator internus." As they
bear upon what I have just said in the Last paragraph,
they will be of value for comparison in the present con-
nection, and so are here republished : this applies more
particularly to the figure (53 his), also given, as it repre-
sents the parts in question as they occur in that curious
bird the Pinon Jay, one of the Corvidce of the western

Fig. 53 bit. — Right three-quartering view of the pelvis of a specimen of the
Pinon Jay (Gyanocephalus cuanocrphalus), showing the origin of the
obturator vntcrnus muscle of the right side, o.i. Drawn by the author
from the specimen, and somewhat enlarged.

part of the United States. In the paper to which I
have just referred, I said, substantially, of the obturator
internus, that " In birds this muscle arises, as shown in
the figure, from the ventral surface of the pelvis, its
fibres being attached to the post-pubic bone and the
ischium. As a rule it is a bipenniform muscle, its fibres
being directed forwards, but at the same time, on either

THE MUSCLES OF THE LOWER EXTREMITY. 193

side of its own moiety, towards a longitudinal tendinous
and mid-line of its own. This tendon becomes stronsrer as
it approaches the obturator foramen, and passing through
this, is finally inserted into the head of the femur of
the corresponding side, and upon its outer surface.

" Now in a great many birds the area from which the
obturator internus arises is of an oval outline, while on
the other hand in nearly an equal number of the class, this
area will be found to be a triangular figure. So it has
been said, that it can thus be utilized as a good charac-
ter, in this way, when taken in connection with others.
In some few birds, I understand, it is difficult to deter-
mine whether this area of origin is oval or triangular,
but as a rule no such difficulty presents itself. For my
own part it constitutes a difference which, I am free to
confess, I had as yet paid but little attention to, as for
several years past neither the proper material nor other
facilities for such investigations have been available.
Nor am I quite sure in my own mind as yet, how far the
form of the hinder portion of the pelvis may influence
the origin of this muscle ; and whether such birds do
not exist wherein a large obturator internus is demanded,
and where their pelves are short, in which cases the
muscle, to gain a firmer origin, would naturally spread out
posteriorly, and thus of necessity become triangular.

" But as I say, I am not prepared to pass final judg-
ment on this matter, and render a personal opinion as
to whether much reliance can be placed upoa it as a
useful character in determinino; affinities amons; birds.

" Here then again is a field open to decide an important
point, and one easily to be understood, and not difficult
to render extensive records about. Those living where
land and water birds occur in abundance could soon
determine whether (or no) this character possessed any

0

194 THE MYOLOGY OF THE RAVEN.

taxonomic value or not, and the result would surely prove
of service to ornithology."

106. The (jtmeUus^ is a strong, thick, rather chunky
muscle, ensconced between the posterior aspect of the
femoral trochanter and the pelvis. It arises from the
entire base of that circumscribed fossa found between the
acetabulum and the obturator foramen, on the outer side
of the pelvis (Fig. 24). Its fibres, attached by fascia to
the tendon of the obturator inter mis, pass directly to
the trochanter of the femur to be co-inserted with the
last-named muscle, to wdiich it plays really the part of
an auxiliary. Professor Owen found this muscle in
Apteryx as a single, small, fleshy strip, and Mivart
says that in some Vertebrates it may be wanting
altogether, as it is in the Ornithorli3aichus and Echidna
{Elem. Anat., p. 342).

The two obturator muscles and the present one are
devoted to drawing the pelvis forward, and steadying
it on the head of the femur when that bone is fixed.

107. The adductor lougus^ is a broad, fiat muscle,

^ Watson followed Owen in regarding the present muscle as the
gemellus ; while others have considered it to be the ohfio-afor ex-
ternus. The subjoined synonymy is from Gadow {loc. cit., p. 173) : —

" 42. Mm. accessorii m. obtueatoris.
L' accessoire de Viliaque interne. Vicq d'Azyr, p. 273, No. 9.
Ohne Namen erwahnt. Meckel, Archiv, p. 266, No. 13 ; System, p.

350, bei No. 10.
Gemellus. Owen, Apteryx, p. 292.

„ Kuhl, Beitrdge, p. 79 ; Watson, p. 108.

Obturator externus. Quennerstedt, p. 15.

,, „ Neander, p. 12.

„ „ De Man, p. 127, No. 5.

Pyramidal {\). Alix, p. 433,
Mm. accessorii tn. ohturatoris. Gadow, No. 20."

2 So distinct ai-e the adductor muscles in the thigh of a Raven,
that I felt myself to be correct in describing them as two. Very

THE xMUSCLES OF THE LOWER EXTREMITY. 195

comjDOsed of rather coarse carneous fibres. It arises
from a line on the lateral aspect of the pelvis, which
line constitutes the lower boundary of the ischiatic
fossa that contains the ohturator extermis muscle. The
extent occupied on this line by the adductor longus
is equal to the length of it between the obturator
foramen and a point rather posterior to the ischiatic
foramen, in the vertical line. Here it is met by the
adductor magnus, the anterior point of its origin. By
an oversight in Fig. 24, the relative positions of the
orio-ins of the semitendinosus and the two adductors are
in each case a little too far forward to agree with what
we find in the majority of specimens. This has been
corrected, however, in the figures illustrating these
muscles, and in the several descriptions.

generally, however, they have been considei-ed as one muscle, and
Gadow records the following synonymy for them {loc. cit., p. 174) : —

" 43. M. PUB.-ISCHIO-FEMORALIS.

M. Septimus femoris. Steno.

Le jpremier adducteur de la cuisse. Vicq d'Azyr, p. 278, No. 2

(nicht le deuxi^me, wie Tiedemann angiebt).
Kurzer Lendenmuskel. Merrem, p. 158, No. 4.
Abductor secundtis femoris. Wiedemann, p. 97.

,, „ „ Tiedemann, § 291 (= adductor magnus

hominis).
Unterer ausserer + innerer Anzieher. Meckel, System, p. 358, No. 8 u. 9.
Anzieher. Meckel, Archiv, p. 264, No. 9.

Abducteurs (pt.). Cuvier, p. 506 ; Gervais et Alix, p. 31 ; Alix, p. 435.
Adductor internus et externus. d' Alton, p. 33.
Adductor femoris longus et add. f em. magnus. Gurlt, p. 28.
Adductor magnus. Owen, Aj^teryx, p. 292.

„ „ Selenka, p. 141, No. 80.

De Man, p. 123, No. 7.

,, ,, Watson, p. 106.

Adductor magmis {et brevis). Quennerstedt, p. 20.

,, ,, ,, Neander, p. 14.

M. jndw-ischiofemoralis. Gadow, No. 18."

0 2

196

THE MYOLOGY OF THE RAVEN.

From the origin 1 liave just given for the adductor
longus, its fibres pass downwards and forwards to be
inserted on a longitudinal line adown the posterior

''o- <ui.uxlal

Fig. 54. — The pelvis of a Kaven, seen upon its ventral aspect, with the skeleton
of the tail. Shows the area of origin of the ohturator internus muscle,
as well as the muscle itself. The origin of the left fcmoro- caudal is
also shown. Life-size, by the author, from his own dissections.

aspect of the shaft of the femur (Fig. 46), from a point
rather above the insertion oiiheJ'e7no7'o-caudal, all the
way to the base of the internal condyle of that bone.
The posterior margin of this muscle is connected to

THE MUSCLES OF THE LOWER EXTREMITY. 197

the anterior margin of the adductor magnus by a firm
but delicate fascia.

108. The adductor magnus is in reality a longer
and narrower muscle than the preceding, and is com-
]30sed of finer and somewhat more compact muscular
fasciculse. It also arises from the line constitutino; the
lower boundary of the ischiatic fossa for the obturator
externus muscle, on the outer lateral aspect of the pelvis.
Its extent of origin on this line commences where the ad-
ductor longus terminates posteriorly, and occupies rather
more than seven millimetres of it in the posterior direction,
to a point where the semitendinosus terminates anteriorly.

From this origin its fibres pass directly down to the
superior curve of the internal femoral condyle, where
they are inserted. The internal head of the gastro-
cnemius is largely attached to this muscle just above
its insertion ; its anterior margin is also attached ]:)y
fascia, as alluded to above, to the adductor longus.
In fact l:)oth of these adductors lie practically in the
same plane, and act as one muscle. We observe also
that they develop but very little tendon, either at
their orio;ins or their insertions.

Owen found the adductor magnus in the Apteryx to
be " a broad and flat muscle, which has an extensive
origin (two inches) from the outer edge of the ischium
and the obturator fascia ; its fibres slightly diverge as
they pass downward to be inserted into the back part
of the lower half of the femur, and into the upper and
back part of the tibia" (loc. cit., p. 101).^

^ There ai-e a nnmber of muscles described by Sir Richard Owen
for the Apteryx that from one reason or another I have thus far
failed to identify with any of those I found in the Raven.

Of these we are to notice the iliacus internus, the pyramidalis,
the adductor hrevis femoris, and the quadratus.

According to this authority, the iliacus internus in the Apteryx

198 THE MYOLOGY OF THE RAVEN.

Referring to the myology of Geococcijx californianus
and the four hist muscles we have just been considering,
we find that the obturator internus arises from an oval
area, and has much the same origin and insertion as
we find it here in the Raven. In (jleococcyx, too, the
gemellus (Fig. 65 his) is a short, thick, carneous muscle,
which arises about the outer rim of the obturator foramen
of the pelvis. Its fibres passing obliquely upwards and
forwards are inserted with the tendon of the obturator
internus muscle on the trochanter of the femur. This
bird also has a few of the fibres of its gemelhis muscle
inserted into the tendon of the obturator externus muscle,

" is a somewhat short thick muscle, of a pavallelogrammic form,
fleshy throughout ; rising from the tiiberosity of the inuominatum
in front of the acetabulum immediately below the gluteus minimus,
and inserted at a point corresponding to the inner trochanter, into
the inner side of the femur near the head of that bone, which it
thus adducts and rotates outwards. This muscle is present both in
the Ostrich and Bustard.

" The pyramidalis arises fleshy from the outer surface of the
ischium for the extent of an inch, and converges to a broad flat
tendon, which is inserted into the trochanter femoris, opposite, but
close to, the tendon of the gluteus minimus, which it opposes, ab-
ducting and rotating the femur outwards.

" The adductor hrevis femoris arises from the innominatum
immediately behind the acetabulum, passes over the back part of
the great trochanter, becomes partially tendinous, and is inserted
into the back part of the femur.

"The quadratus is a broad fleshy muscle which arises from the
pubis, below the obturator foramen, and which increases in breadth
to be inserted into the femur internal and jjosterior to the obturator
tendon " {Anat. of Verts., vol. ii. pp. 100-102).

Since writing the above notes, I have gone more fully into the
literature of the myology of birds, made possible by better
facilities, and it will be evident from the synonymy I have added
from Gadow throughout this work what some of these muscles of
Owen's are ; as, for instance, the pyramidalis is evidently my
obturator externus ; or ISTo. 40 of Gadow, his m. ischiofenioralis. —
R. W. 8. (June 25, 1889).

THE MUSCLES OF THE LOWER EXTREMITY. 199

at least I found this to be tlie case in at least one specimen
examined by me.

The adductors arise from the infero-external margin
of the ischium, between the anterior edge of the semi-
memhranosus and the obturator foramen.

The adductor longus (Fig. 64 bis) is the more anterior
of the two, and consequently arises the higher on the
pelvis, and comes off in front of the adductor magnus,
which it largely overlaps. Its fibres pass obliquely to the
posterior aspect of the shaft of the femur, down which
they become inserted as far as its middle, along the linea
aspera, a line which is well marked in our subject.

The adductor magnus (Fig. 64 his), like the one just
described, is also a broad ribbon-like muscle, arising
from the ischium between the sonimembranosus and a
middle point on the under side of the adductor longus,
close up to its semitendinous origin. Anteriorly its
margin is free, while posteriorly it is juxtaposed to
the anterior border of the semimemhranosus. Passing
parallel with those of the other adductor, its fibres are
inserted into the distal moiety of the linea aspera of
the femoral shaft, down to the intercondyloid notch of
that bone, where this muscle makes a very substantial
insertion.

Other authorities at my hand have but little to say
about the adductor muscles in the thighs of birds.

MUSCULATURE OF THE LEG AND THE REMAINING PARTS
OF THE LOWER EXTREMITY.

The preparatory dissection necessary to bring the
muscles of these parts into view has already been sug-
gested above.

200 THE MYOLOGY OF THE RAVEN.

109. The gastrocnemius'^ muscle in the Eaven is
quite a complicated one, more complicated even than we
find it to be in a man, or others of the higher ty})es of
Mammalia. It develops three distinct heads, viz. an
external head, an internal head, and a tiljial head.

The external head arises rather far Ijack upon the
external aspect of the outer condyle of the femur, Ijy
a short, somewhat flattened, though a strong tendon.
This oriscin is below the orio-in of the lono; end of the
fibrous loops for the biceps, while the extremity of the
short end of this loop is attached to the tendon of
the external head of the gastrocnemius first, before it
passes into its carneous portion. In form this muscular
portion of the external division of the gastrocnemius is
a broad flattened spindle ; slightly concaved on its inner
surfcxce, and rather more convexed on its outer aspect.
The lower apex of this portion of the muscle merges into

^ " 49. M. GASTROCNEMIUS.

31. primus 2Wsterior pedeni et digitos movens. Aldrovaudi.

M. sejitimus circa tibiam et fibulam. 8teno.

Les muscles jumeaux. Vicq d'Azyr, p. 283, No. 1.

Grosser Wadenmuskel. Merrem, p. 4G0, No. 3. »

Gastrocnemius. Wiedemann, p. 101.

,, Tiedemann, § 304.

,, Quennerstedt, p. 32.

,, Neander, p. 20.

DeMan, p. 129, No. 17.

,, Gadovv, No. 33.

,, WatsoD, p. 116.

Les gastrocnemiens. Cuvier, p. 539.
Wadenmuskel oder Fussstrecker, Meckel, System, p. 373, No. 3 ;

Archiv, p. 273, No. 3.
Gastrocnemius internus + externus. Owen, Apteryx, pp. 294, 295.
Gastrocnemien et soleaire tibial. Gervais et Alix, pp. 34, 35.
Gastrocnemien (Jumeau externe et interne + soleaire tibial). Alix,
p. 451." (Gadow, loc. cit., p. 183.)

THE MUSCLES OF THE LOWER EXTREMITY. 201

tlie broad tendinous expansion, about opposite the junction
of the lower and middle thirds of the tibial shaft.

The internal head arises from the outer surface
of the inner condyle of the femur, and rather more
posteriorly in point of situation than the corresponding
origin of the external head. It is broad and more

Fig.Sf.

Fi^. S9.

Fig. 55. — Anterior view of tarso-metatarsus of a Raven.

Fig. ■^^ — Posterior view of tlie same bone.

Fig. #7. — The summit of the same seen from above.

Fig. 58.— Basal joint of hallux, seen from above ; the joint taken from the same

foot.
Fig. 59. — The same bone seen fi'om beneath.

In the figures of the tarso-metatarsus the accessory or liallux metatarsal bono
is in situ. Drawings designed to show tlie origin and insertion of muscles ; and
all life-size, by the author, from his own dissections.

fleshy in character, while the distal end of the adductor
niagnus muscle makes a tendinous connection with the
outer edge of this head, close to its origin, in a manner
already described above. This internal, or what is really,
more correctly speaking, the middle division of the
gastrocnemius is the smallest by all odds. Its fibres
pass directly down the middle of the back of the

202 THE MYOLOGY OF THE RAVEN.

leg, and merge into the l)rocid tendinous expansion
already alluded to in tlie last paragraph. The free edges
of this portion are firmly attached by strong fasciae
to the mesial margins of the other two divisions. The
tendon of the biceps passes between this head and the
external one, while the fascia of the accessory scmitcn-
dlnosus is attached to its outer free edge above.

The tibial head of the gastrocnemius has a l)road
and fleshy origin from the entire inner rim bounding
the tibial summit and from the free edge of the adjacent
procnemial crest (Fig. 50).

This division of the muscle is somewhat laro;er than
the external one, though of a very similar form, its
fibres below converge into a point which is situated
rather lower down than the distal terminations of the
carneous portions of either of the other divisions, and
this point merges into the narrower portion of the
common tendon of the muscle. This tendon, the super-
ficial and median one of the lower fourth of the leg,
develops in it, between the distal tibial condyles, a
semilunar flattened piece of cartilage, which rides in
a longitudinal groove over the true tibial cartilage,
and is braced into position by a firm fascia, at the
back part of the tarsal joint. Below this, the tendon
of the gastrocnemius, hardly reduced in size, though
gradually becoming thinner, passes down directly
over the other tendons at the back of the tarso-
metatarsus, to merge into the fascia over them at
about the middle of the l)one. In all the lower part
of its course it is held in its position by a strong
fibro-tendinous fascia, fully capable of resisting the
most powerful contractions of the muscle.

By way of comparison with what we have just
given above for the Raven, we find this muscle wonder-

THE MUSCLES OF THE LOWER EXTREMITY. 203

fully well developed in Geococcyx. All three of its
heads are strongly defined, and the fleshy belly of the
muscle is massive and thick.

Its external head arises, curiously enough, by two
perfectly distinct tendinous slips. One of these, a strong,
flat tendon, comes off from the outer surface of the
external condyle of the femur, while the second slip, also
strong but somewhat more rounded, arises from the back
of the external femoral condyle, just above the trochlear
surface. Between these two tendons of the external
head of the gastrocnemius we find the loop iov ih.Qhiceps
and the tendon of that muscle itself, the loop being cjuite
intimately attached to the free edge of the outer tendoiL
Below the loop, these tendons merge with each other
and terminate in the commencing fibres that compose
the external head of the gastrocnemius proper.

The internal head of the gastrocneuiius, or what is
really the middle head in birds, is quite median in
position, and is represented merely by a long, narrow,
muscular slip that arises by a delicate, though strong,
cord-like, tendon from the middle of the intercondyloicl
notch of the femur.

The tibial head of the muscle under consideration
is massive in its dimensions when compared with the
divisions of origin of the gastrocnemius already described.
It arises fleshy from an extensive surface on the inner
aspect of the head of the tibia as high up as the marginal
boundary of its summit ; and from the muscular fascia
surrounding certain of the deep thigh-muscles, which are
inserted into the distal end of the femur, and conse-
quently are adjacent to the posterior aspect of the head
of the tibia.

At a point about opposite the junction of the upper
and middle third of the shaft of the tibia the internal

204 " THE MYOLOGY OF THE RAVEN.

and tibial heads of the gastrocnemius mero^e with each
other, while between their free edges above passes the
exceedingly delicate tendon of the semimemljranosus
muscle.

All of the fibres of this complicated origin of the gastro-
cnemius muscle now converge and pass directly down
the back of the leg of the bird. They also merge with
each other in such a manner that, were we to examine
the muscle at about the middle third of the leg, we should
find it composed of two well-defined bellies, rather thin,
nearly of ecj[ual size, united somewhat firmly by an
intervening fascia, and each being convex on their
superficial aspect and the reverse on their under sides,
which concavity accurately moulds itself to the deeper
layer of muscles of the leg, which the gastrocnemius
completely covers.

At the lower fourth of the tibial shaft the fibres
terminate in a broad, flat, and glistening tendon, which
passes flat- wise over the shallow and longitudinal groove
of the tibial cartilage, at which point the tendon is
considerably thickened. Next, crossing the tibio-tarsal
joint, it becomes internally attached to the hinder surface
of the hypotarsus of the metatarsal bone, below which
protuberance it finally merges into the deeper layer of
the podothecal sheath confining the flexor tendons.

110. Tlte soleus^ is exposed when we remove
the tibial division of the gastrocnemius. It is found

^ Provisionally, at least, I still propose to regard this muscle as
the soleus, being by no means yet satisfied that it can be considered
the homologue of the 2^l<^nta'ris of the Mammalia. Aside from
everything else, however, attention is especially invited to the fact
that in the vast majority of birds it arises from the tihia, and its
tendon below merges with fhe gastrocnemius. In support of its being
the homologue of the plantaris, Gadow has said : — •

" Bei einigen Yugeln entspringt der Muskel waiter proximalwiirts.

THE MUSCLES OF THE LOWER EXTREMITY. 205

to be a somewhat flattened, small muscle, pointed
below, but arising by carneous fibres from quite a
broad base at the back of the head of the tibia. The
broad, thin tendon of insertion of the semimembran-
osus overlaps its belly from the inner side. From its
apex below it sends down a long, slender tendon,
which is inserted into the proximal end of the
" tibial cartilage," towards its inner angle. In the
Apteryx, Owen found that the tendon of this muscle

namlich von der Hinterfliiche des Gondylus interims femoris, doch
kann dies durchaus nicht so hiiufig sein als Meckel angiebt, denn
ich fond Ein femoralen Ursprung nur sehi' selten, z. B. bei
Corythaix, bestiitigt. . . .

" Entspringt der M. 2^lot'ntar{s vora Femur, obgleicli vom inneren
Condylus, so ist er beinahe vollstandig dem gleichnamigen Muskel
des Menschen homolog. Bei den Reptilien ist ein solcher Muskel
in dem noch ungetheilten Flex. long. dig. enthalten."

I by no means consider this argument as being especially cogent
in support of the muscle being considered the jilantaris. Far more
extended observations upon the myology of the Vertebrata generally
ax^e required to decide such a point as this. Gadow gives further
the following synonymy {loc. cit., pp. 185, 186) : —

" 50. M. PLANTARIS.

La grele plantaire. Vicqd'Azyr, 283, No. 2.

Plantaire. Cuvier, 539.

Flantaris, vielleicht hinterer Schienbeinmuskel. Meckel, System,

375, No. 4.
Ohne Namen. Reid, pp. 144, 145.
Soleus. Owen, 295.
Jamhier 2>osterieur. Gervais et Alix, 35.

Alix, 452.
Flantaris. De Man, 130, No. 18.

,, Gadow, No. 40.

„ Watson, 119."

JVote. — I observe that Selenka, in his drawing of the muscles of
the lower extremity of a Gull (Larus/uscus), marks this muscle as
the soleus (see Bronn's Klassen des Thier-Reichs, vi. Band, Taf.
xxiii).— R. W. S.

20G

THE MYOLOGY OF THE RAVEN.

"joins that of the gastrocnemius internus, Lehiiid the
tarsal joint" (Anat. of Verts., vol, ii, p. 106).

The soleiis (Fig. 65 his) is a well-developed muscle in
Geococcyx calif or nianus. It arises from behind the

Fig. 60. —The skeleton of left foot of a Raven seen from behind ; designed to
show origin and insertion of muscles, and drawn life-size, by the author,
from his own dissections. a, line of tendon of the Jlcxor pcrforatus
medius prh/uos pedis, b, is the line of the tendon of the flexor perfora-
tits annularis primiis pedis. The exact insertion of these tendons are
somewhat hidden by the hind claw and basal joint, and they are drawn
to some extent diagrammatically.

tibia, on its inner side, and just below the marginal rim
of its summit. The fibres at once form a little flat
muscle, rather longer in shape than the fish from which

Fig. 61. — Outer aspect of the left pelvic
limb of a Raven, showing the
siipeihcial nmscles of the thigh
and leg. Life size, by the
author, from his own dissec-
tions.

208 THE MYOLOGY OF THE RAVEN.

it derives its name, and soon terminate at the lower or
tail-end in a tendon. This tendon, long and narrow,
passes directly down the postero-internal aspect of the
leg to become inserted into the dense fascia covering the
tibial cartilage at its supero-internal angle.

Professor Mivart says that in Ornithorhynchus this
muscle is inserted into the astragalus. And that fur-
ther, in Nycticehus it has lost its tibial attachment,
is entirely muscular, and blends with the gastrocnemius.
This same author also informs us that in the Agouti
it arises from the tibia only, wdiile it is wanting in the
Pig, Hyaena, Seal, and others {Elem. Anat., p. 354).
I present these comparative notices of such muscles
as the soleus in others of the Vertebrata in order that
we may have before us at least short histories of the
muscle in question, which often prove suggestive.

Another word here in regard to the "tibial cartilage."
I find a number of specimens, apparently very old
birds, wherein its lower outer angle has ossified, and
c[uite a sizable sesamoid is there formed, wdth a facet
for the tibia.

111. The peroneus longus^ is very well developed

^ A very full description of this muscle is given by Professor
Gadow, to whom we are also indebted for the subjoined nomen-
clatural record, to wit : —

" 47. M. PERONEUS SUPERFICIALIS.

M. quartus posterior p)^deni movens. Aldrovandi.

M. decimus circa tibiani et fibulam. Hteno.

L" accessoire des flechisseurs des doigts du pied. Vicq d'Azyr, 1774,

p. 510, No. 1.
Innerer Beinmuskel. Merrem, p. 160, No. 2.
Extensor tarsi externus. Wiedemann, p. 99.

M. tibialis posticus. Tiedemann, § 305 ; Carus, Erlauterungstafeln.
LaDger oberer Wadenbeinmuskel. Meckel, S)jstem, p. 384 (innerer

stiirkerer Kopf von No. 2).

THE MUSCLES OF THE LOWER EXTREMITY. 209

in our present subject, being a broad, muscular sheet,
that covers or nearly conceals from view all the muscles
on the anterior aspect of the leg, and principally the
tibialis anticus.

It arises from the raised cnemial crest in front of the
head of the tibia, and from the fascia that covers the
outer side of the knee-joint. The fibres, forming a closely-
fitting, muscular curtain eml)racing the frout of the
leg, pass downwards, then downwards and outwards to
terminate in a small tendon at about the lower third of
the tibia. This tendon bifurcates just above the tibial
condyles at the outer aspect of the limb. The shorter
and stronger fork of the bifurcation goes to the upper
end of the tihicd cartilage, to become attached to the
fibrous fascia covering it ; while the smaller slip passes
to the outer side of the hypotarsus of the tarso-meta-
tarsus, to meet the tendon of the Jiexoi' j^erforatus medius
2:>rimus pedis obliquely, and merge with it at the back
of that bone, about one centimetre below the hypotarsus.

Langsehniger Sohlenmuskel. Meckel, Archiv, p. 273, No. 4.

Moyen peronier. Arvier, p. 542.

Peroneus medius. Owen, Comp. Anat., ii. p. 108.

Soleus et peroneus longus. Gurlt, p. 30.

Feroneus longus. Owen, Cyclopctalia, p. 297 ; Apteryx, p. 296.

,, ,, Quennerstiidt, p. 36.

,, ,, ISTeander, p. 21.

De Man, p. 134, No. 24.
Watson, p. 123.
Peroneus longus s. communicans. Nitzsch, in Giebel's Zeitsclirift,

X., 1857, p. 24 u. 240.
Long 2)ero7iier. Gervais et Alix, p. 34 ; Alix, p. 450.
M. pieroneus superjicialis. Gadow, No. 29."

Note. — I question whether the peronexis medius of Owen (Comj).
Anat., ii. p. 108) is the present muscle as is qvioted in the above
synonymy ; but am inclined to think that the j^^foneus longus of the
same author, on p. 107 of the same woik, may more properly be
considered so. — E.. W. S.

P

210

THE MYOLOGY OF THE RAVEN.

I designate as the tibial cartilage that fibro-carti-
laginous block lying between the tibial condyles
behind, over which the tendon of the gastrocnemius
passes, and through which the tendons of the flexors
of the podal digits glide. Thus we see that the peroneus

Fig. 62. — The skeleton of the left foot of a Kaven, seen from the side ; designed
to show passage and insertion of tendons, &c. Drawn life-size, by the
author, from his own dissections.

longus in the Raven takes the part of an auxiliary to the
flexors of the toes, assisting them in their action.

According to Sir Richard Owen, the present muscle in
the Apteryx behaves in a very similar manner in the
lower part of the leg to what I have just described

THE MUSCLES OF THE LOWER EXTREMITY. 211

for the Raven. We find in tlie Apteryx that the
" peroneus longus arises tendinous from the head of
the tibia, and by carneous fibres from the upper half
of the anterior margin of the tibia ; these fibres pass
obliquely to a marginal tendon, which becomes stronger
and of a rounded form where it leaves the muscle.
The tendon gives off a broad, thin, aponeurotic sheath,
to be inserted into the capsule of the tarsal joint; it
is then continued through a synovial pulley on the
side of the outer malleolus, and is finally inserted or
continued into the perforated tendon of the middle
toe" (Aiiat. of Verts., vol ii. pp. 107, 108).

In reflecting the peroneus longus in the Raven, and
viewing its under side, we observe that where it takes
origin from the free points of the pro- and ectocnemial
processes it is very tendinous, and these tendons can
be distinctly traced down on this surface of the muscle
for some little distance, forming lateral boundaries to
the more carneous central portion.

112. The tibialis anticus ^ is a very interesting muscle

^ This muscle has been very generally considered to be the
tibialis anticus by morphologists, as will at once be appreciated by
the synonymy given us by Gadow, viz : — -

" 45. M. TIBIALIS ANTICUS.

M. secundus anterior jy&deni movens. Aldrovandi.
M. undecimus circa tibiam et fibulam. Steno.
Le tibial anterieur. Vicq d'Azyr, 1774, p. 510, No. 2.
Anzieher des Fusses. Merrem, p. 164, No. 4.
Tibialis anticus. Wiedemann, p. 99.

,, ,, Tiedemann, § 306.

„ „ Owen.

,, ,, Neander, p. 22.

,, ,, Quennerstedt, p. 38.

De Man, p. 135, No. 26.

„ „ Gadow, No. 31.

„ ,, Watson, p. 48.

Le tibial ou jambier anterieur. Cuvier, p. 539 ; Alix, p. 448.

P 2

212 THE MYOLOGY OF THE EAYEN.

in the bird we now liave under consideration. Its
carneous portion is divided into two very distinct parts,
which, though moulded together, are easUy separable
clear down to that point where they unite with their
common tendon below. The inner head^ — if we may be
permitted to so term these divisions— arises immediately
beneath the peroneus longus from a semilunar area
high up between the pro- and ectocnemial crests on
the anterior aspect of the head of the tibia. This
division forms the anterior half of the muscle, its origin
being quite extensive, and principally fleshy. The
" outer head " of the muscle arises by a short, strong
tendon from the base of a little pit found upon the
antero-inferior ridge of the outer condyle of the femur.
This head is overlapped by the fascia of the knee-
joint and the great flexor at the 1)ack of the leg, but,
passing beneath these, it immediately moulds itself
upon the anterior half, though there is no blending of
fibres whatever, it being situate completely behind it.
The two then pass directly down the front of the leg
as a large and handsome fusiform muscle. At the lower
third of the bone their fibres converge to a point, to
become attached to a strong and powerful tendon
common to the two divisions ; this, passing through the
oblique, fil)ro-cartilaginous bridge just above the tibial
condyles, goes directly, in the antero-median line, to
a point on the shaft of the tarso-metatarsus just below
the head of that bone, where it is finally inserted upon
a tul^ercle, there found, and which is intended for it.

Both the j^c^'onetis longus and the tibialis anticus are
well developed in such a bird as Geococcyx calif or nianus,
and have essentially the same origins and insertions as

Vorderer Schienbeinmuskel. Meckel, System, 370, No. 1 ; Archiv,

p. 272, No. 1.
Levator 2)edis. d' Alton, p. 36."

Fig. G2 his. — Outer aspect of the riglit pelvic limb of the "Eoad Painnei"
(Geococcyx calif or nianus), showing the superficial layer of muscles, and
the relations of the peroncus longus and the tibialis anticus are especially
to he observed. Life-size, by the author, from his own dissections.

214 THE MYOLOGY OF THE EAVEN.

we have described for them above, as they exist in the
Raven. We have good views of them in Fig. 62 bis of
the present work. Several years ago I devoted no little
attention to the myology of Geococcyx, and published
my observations thereon in a number of places. The
bird, as is well known, occurs on the Pacific coast region
of the United States, and presents much of interest in
its anatomy. It is a big Ground Cuckoo with affinities
in other directions.

To become impressed w-ith the gTcat numljcr of
changes to which this muscle is subject, we can do no
better than to read Professor Mivart's account of it, as w' e
find it among other representatives of the Vertel:)rata.

This eminent anatomist tells us that " the tihialis
anticus, even in Anthropoid Apes, may have the part
ffoing to the hallux so distinct as to l)e reckoned a
distinct muscle— sometimes called the abductor longus
haUucis. It may be uncpiestionably double, as in the
Echidna. It may have a double origin and single
insertion, as in the Agouti. It may be inserted into
the second metatarsal, as in Hyrax ; and may be
altogether wanting, as in the Pig. It is inserted into
the tarso-metatarsal bone in Ijirds, and is situate
quite on the inner aspect of the leg in Cliameleo. It
exists down to the Urodeles, being apparently doul)le
even in Menobranchns. It may arise from the femur,
as in the Frog" (Elem. Anat., p. 351).

Owen also found the tibialis anticus overlapped
by the peroneus, and arising partly in common with
that muscle, " and partly by sejiarate short tendinous
threads from the outer part of the head of the tibia ; it
gradually becomes narrower, and finally tendinous tw^o-
thirds of the way down the leg ; its strong tendon
glides through the oblique pulley in front of the distal
end of the tibia, expands as it passes over fhe ankle-

THE MUSCLES OF THE LOWER EXTREMITY. 215

joint, and is inserted into tlie anterior part of tlie
proximal end of the tarso-metatarsal bone, sending off
a small tendinous slip to the aponeurosis covering the
extensor tendons of the toes, and a strong tendon
which joins the fibular side of the tendon of the ex-
tensor longiis digitorum'' {Anat. of Verts., vol. ii. p. 108),

In man, we remember, the tibialis anticus is a direct
flexor of the tarsus upon the leg.

113. The extensor longiis digito^'um'^ (Fig. 51) arises
fleshy from the under edge of the procnemial ridge or
crest of the tiliia, its mesial side, and from a limited
area of the contiguous surface of the shaft of the l;)one.
Its fibres pass down on the interno-anterior aspect of
the tibial shaft, as a long, slender, fusiform muscle.
At the lower third of the bone its strong tendon sup-
phants the carneous portion, and gradually coming to
the middle line, passes under the bony bridge just above
the condyles in front. Emerging from this, it passes
over the front of the ankle-joint, being bound down in
this situation by a firm fascia. It is now slightly

^ A study of the methods of insertion of this muscle throughout
the Class Aves is a very interesting one, and shows it to be very
different in several of the groups. Gadow gives its synonymy as
follows : —

" 46. M. EXTENSOR DIGITORUM COMMUNIS.

M. jyt'imtis anterior 2^eJevi et digit os movens. Aldrovandi.

M. duodecimus circa tibiam et Jibulam. Steno.

Vextenseur commun des doigts. Vicq d'Azyr, p. 282, No. 3.

Schienbeinmuskel. Merrem, p. 161, No. 5.

M. extensor digitorum communis. Wiedemann, p. 100.

,, ,, ,, ,, Tiedemann, § 308.

,, ,, ,, ,, Quennerstedt, p. 39.

,, ,, ,, „ Neander, p. 23 ; Gadow, No. 32.

„ „ „ „ Watson, p. 126.

Extensor digitorum longus. Owen, Aj^teryx, p. 297.

„ „ „ De Man, p. 135, No. 27.

Vextenseur commun des doigts. Gervais et Alix, p. 33 ; Alix;
p. 453."

TvaZZuycis.

Fig. 63. — Outer aspect of left pelvic limb of a Raven, sliow-
iiig the second layer of muscles. The crurwics
and vasttis extcrn.us are somewhat parted at their
lower moieties, and the biceps drawn a little to the
rear to show its passag;e through the tendinous
loop. Life-size, by the author, from his own
dissections.

THE MUSCLES OF THE LOWER EXTREMITY. 217

deflected to the inner side, and just above the tubercle
on the tarso-metatarsus for the insertion of the tendon
of the tibialis anticus, it passes through a special little
bony arch there found for it. It then passes down the
interno-anterior edge of this bone to the trochlese of
the basal toe-joints. Here it divides into three smaller
tendinous slips ; these pass respectively over the
superior aspects of the second, third, and fourth toes,
bifurcating as they do so, beyond the basal joints, to
be inserted at the bases of the distal ones. At the
pioximal extremity of each series of phalanges, the
tendons are bound down in the median grooves in-
tended for their passage and guidance by a strong,
fibrinous, fascia-like sheath.

This muscle receives on its outer side a separate slip
that Blight almost be reckoned as a minor head were it
not in such a rudimentary state.

It aiises as a small bundle of carneous fibres from
a limited elliptical area on the front of the shaft of
the tibia, near its middle, and about opposite the prox-
imal end \)f the fibular ridge. It passes down to the
outer side t>f the extensor longus digitoriim proper, and
in close contact with it, and about two centimetres down,
it develops \ feeble, fascia-like tendon, which with the
terminal fibre!\^of the muscle itself soon merge with those
of the large ext?,nsor at its side. The tibialis anticus must
be removed bef<a'e this delicate little slip can be brought
into view, as it ies immediately beneath it (Fig. 51).

Upon another pccasion, when speaking of this muscle
as it is found in Geococcyx, I said : —

"The extensor l6\gus digitorum (Fig. 64 his) arises from
the anterior aspect ^ the inner half of the tibial shaft
as high up as the tibi^is anticus muscle, which covers it ;
it also arises from a i^nse fascia which comes off from
the lower free edge of \the procnemial crest of the tibia ;

218 THE MY0L0(;Y OF THE RAVEN.

and finally from a longitudinal line extending obliquely
down tlie front of tlie shaft of tlie tibia to its lower third.
This oljliquity finally brings the tendon in wdiich the
extensor long us d ig itorum tenoninates, to the middle line.

" Just above the condyles of the tibia, it here passes
through the little bony bridge ; emerging from which it
crosses the ankle-joint in front, then passes down the
anterior aspect of the tarso-metatarsus bone, overlying
the short extensor. At the upper end of this last-named
bone, and over the ankle-joint, this tendon is firmly
bound down by a fibrous fascia. In some birds we know
a special bony span exists for it on the upper part of the
tarso-metatarsus, as in certain Owds. When the terdon
of this muscle arrives at the anterior aspect of the trochleaB
of the distal end of the tarso-metatarsus, it expands and
bifurcates. The tendinous expansion becomes more or less
attached to the underlying tissues, while each bifi^rcation
passes one over the second, and the other over lhe third
toe, for their entire lengths, to become insertec' into the
upper points of their ungual phalanges.

" Now from the side of the tendon that o^oes :o the third
toe another slip is diff"crentiated ofi" in a V3ry peculiar
manner, owing to the reversion of the toe in question.
For it not only passes over the top lengtl for insertion
of this fourth digit, as in the case of the others, but its
slip also splits ofi" to make a separate insertion at the
extremity of the basal phalanx of th« third digit. I
have had the opportunity of dissectine three feet, with
the view of studying this point, and ^ find it to obtain
in all of them."

114. Tlie extensor hallucis hrevu^ is an exceedingly

^ As I read the above account I h.a e given of the extensor
hallucis hrevis, I am convinced that I \y&> compelled to so name it
myself for our Raven ; and the literaure of the subject was at
that time not available to me, and I 'Jis not aware that it had

THE MUSCLES OF THE LOWER EXTREMITY. 219

interesting little muscle. It arises fleshy from the antero-
internal aspect of the head of the tarso -metatarsus, just
Below its summit ; from the corresponding edge of the
shaft below ; and from the tendon of the tibialis anticus.
The delicate little bundle of fibres pass down the antero-
internal edge of the shaft of the bone, which is slightly
concaved to receive them, in order that the close-fitting
podotheca may not interfere with their proper action ;
when just before arriving at the apex of the accessory
metatarsal, they unite with a delicate, though strong,
little tendon, which, passing round behind that bone
and over the hallucial basal joint, runs along over the
top of the phalanx of the hallux close to the Ijone,
to be finally inserted into the process at the superior
base of its bony claw.

Professor Owen, in the second volume of his Anatomy of
Vertebrates, describes also an extensor brevis digitoruni ;
but that muscle does not occur in the Eaven, and the

received a similar designation at the hands of others. Gadow has
bestowed the same name upon it, and furnished the following
synonymy : —

"54. M. EXTENSOR IIALLUCIS BREVIS.

M. 2»'i'nius anterior tarsi. Aldrovandi.

M. 2^'>'iinus circa os quod siipplet vices ossiiwi tarsi et metatarsi.

Steno.
L'ahducteur du doigt oppose. Vicq d'Azyr (1805), p. 288.
Extensor hallucis. Wiedemann, p. 106.

,, ,, Tiedemann, § 315.

„ „ d'Alton, p. 39 ; Owen, p. 297.

,, ,, Quennerstedt, p. 48.

,, „ Neander, p. 28.

„ ,, Gadow, No. 41.

Vextenseur propre du pouce. Cuvier, p. 553.
Extensor of the thumb. Reid, p. 145.

,, digitorum brevis (pt.). Gurlt, p. 32.

,, brevis hallucis. De Man, p. 138 ; Watson, p. 126.
unguis. Garrod, P.Z.S., 1872, p. 363.
Eextenseur du pouce. Alix, p. 447."

220 THE MYOLOGY OF THE EAVEN.

one I have last described seems to correspond to tliis
writer's extensor pollicis hrevis, but liis description is
not very definite.

I dissected four pairs of feet of this species, — and
the dissection is by no means an easy one, — before I
was satisfied tliat the facts in regard to these short
extensors of the toes are as I have given them above.
They were the same in all, and in all the extensor
hrevis digitorum of Owen w^as absent.

Professor Mivart, in his Eletnentary Anatomy, does not
mention the short extensor of the hallux as occurring
in birds, and this eminent biologist calls this muscle
the extensor proprius haUucis.

In certain birds with zygodactyle feet, as in Geoeoccyx
for instance, the morphology of the extensor hallucis
hrevis is quite different, and when treating the myology
of G. californianus it led me to consider the present
muscle as only a part of the extensor hrevis digitorum
in that form, and I said in effect that the extensor hrevis
digitorum is a muscle that may be taken as an ampli-
fication of the muscle I have described in the Eaven as
the extensor hcdlueis hrevis.

But even here in Geoeoccyx the short extensor of the
hallux has a certain amount of individualization, thousfh
it is not fully differentiated from the other part of this
extensor hrevis (Fig. 64 his). It, however, is not at-
tached more than half-way down the anterior aspect of
the shaft of the tarso-metatarsus, at which point it
terminates in a delicate thread-like tendon ; this passes
directly over the upper border of the accessory meta-
tarsal, and along the top of the basal joint of the hallux,
to become inserted in the usual manner in the base
of the claw-joint. Now the remainder of the extensor
hrevis digitorum is attached down the shaft of the tarso-
metatarsus, as far as the distal trochlese ; the outer

,&*tejvsor fe^nxAivts.

Fig. 63 his. — Pelvic limb, right side, of Geococcyx calif ornianus, same spociineu as
is shown in Fig. 62 his. The superficial layer of muscles have here been
removed, and those found beneath them brought more fully into view.
Life-size, by the author, from his own dissections.

222 THE MYOLOGY OF THE RAVEN.

portion of it developing a tendon about lialf-way down,
which is concealed by the carneous fibres which overlie it.
This tendon passes round beneath the trochlea for the
fourth toe, and is really inserted on the undei'side of the
basal joint of this digit at its proximal extremity ; so
that in the case of this toe it seems as thouo;h it would
act almost as nfiexor. AVith the second and third toes,
however, the carneous fibres of the muscle under con-
sideration are continued all the way to the trochlese,
where they terminate, in either case, in a strong, fiat
tendon, which passing over the joint is inserted on the
U}yper side of the proximal extremity of the basal joint.
Here, of course, the muscle acts (in the case of the second
and third toes) as an auxiliary to the long extensor.

Not a little room is here open to us for speculation as
to how the tendon of this short extensor in the case of
this fourth toe exactly came to assume its present point
for insertion, as the digit gradually and finally became
permanently reversed. Indeed, the high development
of this short extensor in Geococcyx over the vast majority
of the class is, too, an interesting fact ; and did the
reversion of the digit precede or follow the muscular
development ■? No doubt the completeness of the latter,
and its perfection for an avian type, has come about as
a demand on the part of the habits of the bird itself and
its marvellous fieetness of foot.

The muscles of the leg thus far dissected out should
now be carefully examined, particularly in regard to their
relations, and the relations and passage of their tendons.
They should next be removed close up to their origins
and insertions, and all superfluous tissue removed, — as fat,
the vessels and nerves, and fascia and connective tissue — •
leaving the muscles at the back of the limb perfectly
clean, and in as favourable condition to study as possible.
Next the podotheca of the tarso-metatarsus and

THE MUSCLES OF THE LOWER EXTREMITY. 223

toes should be removed with the greatest care, using
a sharjD scalpel for the purpose. This will take fully
an hour, as we should be particularly guarded not to
injure in the slightest degree any of the tendons, or
disturb their mutual relation to each other. At the
end of our work nothing should be left but the
skeleton of the limb ; the muscles and their tendons
in situ ; and the tibial cartilage so trimmed as to show
precisely the manner in which the tendons pass through
it, and their relations to each other.

It is a good plan now to fasten the limb down
to a smooth surface in such a manner that the pos-
terior aspect faces upwards and towards you, then Ijy
means of little hooks and dissecting chains the individual
muscles and their tendons can be raised in such a
manner as to greatly facilitate their study. I am thus
particular in these details because I have experienced
no little difficulty myself in properly comprehending
these parts, and it is simply out of the question to
do it at all hj a hasty dissection. ^v

Now it will be seen, as a sort of a first introduction
to these parts, that the hind toe or first toe is under the
control of two tendons (whatever their muscles may be
above) : the first of these is the delicate little extensor
haUucis hrevis (Fig. 60) already described, and the
second a powerful flexor tendon running along under-
neath its basal joint. This tendon at the back of the
tarso-metatarsus ossifies, and the bone is usually some-
thins; long;cr than two centimetres. Others of the laro-er
tendons in the same situation do the same thing in old
specimens. These bones are shown at the back of the
tarso-metatarsus in Figs. 49 and 60.

Confining ourselves now to the sole of the foot in this
cursory examination, we see that the second toe (with
two joints and a claw) is served by three tendons coming

tiZialis ^ostlctxs

Fig. 64.— Outer aspect of the left pelvic limb of a Raven, showing
the third layer of muscles of the thigh and leer. The
stump of the hiccps has been left in position, and the
horizontal little band of tendon binding the head of
the flexor perforaMos indicis 'primus pedis to the iibula,
as well as affording support to the bicipital loop, is
shown, a = a dissecting chain pulling back the tendon
of the flexor per/oratus annularis primus pedis muscle ;
and b, accomplishing the same for the tendon of the flexor per foratus
mcdius prmius pedis muscle. Life-size, by the autJior, from his own
dissections.

THE MUSCLES OF THE LOWER EXTREMITY. 225

from muscles at the back of the leg. One of these
perforates the other two, and runs the entire length of
the toe, to become inserted at the tubercle at the under
side of the bony core of the claw or the ungual phalanx.
Of the two perforated tendons the longer one is inserted
at the distal end of the , basal phalanx, and the shorter
one at the middle of the shaft of the same bone. In the
middle or third toe (with three joints and a bony claw)
we find also a long perforating tendon inserted as in the
last toe, with the longer perforated tendon inserted at
the distal extremity of the second phalanx, while the
shorter one stops at a point over the shaft of the liasal
joint. Now the outer toe (with four joints and an un-
gual phalanx or bony claw) has but two tendons, one
perforating and the other perforated. The perforating
one behaves as in the other toes, running clear to the
tubercle at the lower base of the ungual phalanx for
insertion, while the perforated one sends down its
bifurcations, one on either side of the perforating tendon,
to be inserted in the one case over the shaft of the basal
joint, and in the other at the distal end of the joint
next beyond, thus making up for the absent tendon in
this toe.

It is evident from this we have in the second and
third toes a perforating tendon and two perforated
tendons, while in the fourth toe we have a perforating
and a perforated one. All of these tendons at the hack
of the tarso-7neta.tarsus hone are separate.

I have carefully read over and compared the writings
upon the plantar or flexor tendons in birds, by Owen,
Garrod, Forbes, Mivart, and Coues, and am convinced
that no little difference of opinion ol:)tains among these
writers, and I regret to say that I am compelled to intro-
duce still another opinion in the case of two or three of

Q

226 THE MYOLOGY OF THE RAVEN.

the muscles iu question, but only after the most con-
scientious study. Nor do I expect my opinion will
hold for all forms of birds, for as I have so often
stated on previous occasions, an enormous amount of
work still remains to clear up even these simple questions
in biology.

Garrod. and Forbes had chiefly to do with a limited
number of the plantar tendons, and I believe, so far as
they go, describe them correctly. Mivart has but little
to say about the musculature of these parts in Aves, and
I fail to find that any of these authorities speak of the
two sets of perforated tendons, with the exception of
Owen, who after descril^ing a flexor 2)e7 for ans digitorum :
ii flexor 2)erforatus of the outer toe ; -a flexor perforatus
digitorum ; and as I say, a 'peroneus longus and medius,
says without any further explanation that " The second
and third toes have two perforated tendons ; one inserted
into the sides of the first, and the other into the sides
of the second phalanx" (Anat. of Verts., vol, ii. p.
109), but says nothing of the muscles from whence the
second set are derived.

In view of the fact that a bird's liand an,d arm
have become completely subordinatcid to the purposes
of flight, and that its foot has become converted largely
into an instrument of prehension, is it not possible
- — nay, quite probable — that a higher specialization
of the muscles has, pari passu, become a necessity ?
and that the peronei muscles (beyond the peroneus
longus with its strikingly feeble insertion) are not
represented in these peculiarly modified vertebrates,
BO far as their extremities are concerned ? These
peronei muscles are by no means constant, and are
chiefly confined to the class Mammals.

Such facts must have their weight in our myological

THE MUSCLES OF THE LOAVER EXTREMITY. 227

nomenclature, and oftentimes, in the present state of
our knowledge of sucli matters, a certainty of diagnosis
in homology is simply out of the question.

In distinguishing the flexors I will bestow such names
upon them as will show their methods of insertion ;
for instance, the ijerforated tendons inserted into the
hasal phalanges will bear the name of primus, with the
name of the particular toe. In this last designation we
will call the second toe indicis, as it corresponds to the
index of manus ; and if occasion calls for it, we will
designate the next toe as medius, and the outside
one as annidaris. It is unnecessary to say that the
term perforans refers to a tendon that 'perfoixUes ; and
perfoi^atus to one that is j^erforated.

Now the perforated tendons of the second row of
phalanges had better be designated by the word secundus,
should they be sufficiently individualized as to warrant
their being described as belonging to separate muscles.

It probably will not be necessary to use all these
names, and I simply throw them out as a suggestion,
though, as I say, if occasion requires it, the nomencla-
ture adopted will be ujDon this j)lan, and in the present
subject some of them will be brought into use.

115. The tibialis posticus arises from the whole
length of the shaft of the fibula below the insertion of
the biceps jiexor cruris ; from the shaft of the tibia
for a little distance below it ; from the interosseous
membrane ; and the adjacent surface of the tibial shaft.
It is in intimate relation with the Jiexor perforans
digitorum pedis, which overlies its mesial portion, and
the two muscles in this situation blending by their
fibres. The tibicdis posticus is a long, subcylindrical
muscle, and passing directly down the leg, from the
origin just described, terminates in a strong tendon

Q 2

228 THE MYOLOGY OF THE RAVEN.

at about the middle of tlie lower tliird of the shaft of
the ti1.)ia. It passes in front of the external malleolus,
crosses the ankle joint, and is finally inserted into the
outer edo-e of the summit of the tarso-metatarsal bone.

This appears to be one of the peronei muscles of
A. Milne -Edwards ; but from the facts that it is evidently
inserted into the tarsus, and not the metatarsus ; and
that the tibialis postimis is quite a constant muscle
among Reptiles ; and that further, as I have already
said, the peronei muscles are more particularly mam-
malian institutions, I believe w^e are safer in calling this
one the name we liave.^

1 The only muscle I can find in literature to compare the pre-
sent one with is the Feroneus profundus of Gadow, who offers the
following upon it (loc. cit., p. 182) : —

" 48. M. PERONEUS PROFUNDUS.

M. tertius anteriur 2^edem inovens. Aldrovandi.

M. tridecimus circa tibiam et fihulmn. 8teno.

Le peronier. Vicq d'Azyr, p. 282, No. 4.

Wadenbeinmuskel. Meckel, System, p. 372, No. 2 ; Archiv, p.

273, No. 2.
Peroneus. Wiedemann, p. 101.

,, Tiedemann, § 307. «•

,, d' Alton, p. 36.

Le court ptronier. Cuvier, p. 542.

„ „ Gervais et Alix, p. 34.

Feroneus tertius. Gurlt, p. 30.

,, brevis. Quennerstedt, p. 37.
,, ,, Neander, p. 22.

De Man, p. 135, No. 25.
,, „ Watson, p. 118.

Peronier lateral. Alix, p. 450.
Peroneus profundus. Gadow, No. 30.

" Der tiefe Wadenbeinmuskel erscheint nach Abtragung des
Peron. superficialis. Er ist ein gewohnlich kleiner, gegliederter
Muskel, der fieischig von der vorderen und ausseren Fliiche der
Perone und von benachbarten Theilen der Tibia, in der Kegel vom

THE MUSCLES OF THE LOWER EXTREMITY. 229

In our Geococcyx callfornianus the tibialis ijosticus
is a veiy slender muscle, but closely resembles tlie same
muscle as I have found it in all other birds which I have
examined for their myology.

As in a number of the Passeres, we find it here to
arise from the aiitero-lateral aspect of the shaft of the
fibula below the tubercle for the insertion of the biceps
Hexor crurisy from the interosseous membrane between
the leg-bones, from the contiguous surface of the shaft of
the tibia, and, finally, from the fascia separating it from
the deep flexors of the leg. The fibres pass directly down
the outer side of the tibia as a long, slender, fusiform
muscle. At the lower fourth of the shaft of this bone
they terminate in a small tendon, wdiich, passing in
front of the external malleolus, crosses the ankle-joint
to become inserted into the supero-external rim of the
summit of the tarso-metatarsus.

116. The jiexoT perforatus indicis secundis pedis ^ is

vorletzten Yiertel des Untersclienkels entspringt. Die AusdelmuDg
des Ursprunges variirt jedocli ungemein.

" Insertion. Der Muskel gelit in eine starke, rundliche Sehne
iiber, welche am aussereu Malleolus durch ein Retinaculum 2^6ronei
tritt und sich dann an der proximalen Ecke des mittleren oder
hintersten der drei Tarso-Metatarsalknochen inserirt."

Note. — These remarks are here added on June 25, 1889, and in
concluding I would say that the muscle certainly has an extensive
fibular attachment in Corvus, and although it may eventually
prove to be a peroneus muscle, I was in no ways influenced in
naming it by any semblance it might have to that muscle so called
in Homo, but rather by the fact that a tibialis posticus occurs in
certain reptilian forms. — R. W. S.

^ This muscle has the following account of its synonymy given by
Professor Gadow : —

" 52a. M. FLEXOR PERFORANS ET PERFORATUS DIGITI IL

M. fertius j)osterioT circum tibiam et fibidam. Steno.

230 THE MYOLOGY OF THE RAVEN".

in the Eaven a perfectly distinct muscle, and fully merits
a separate description. It arises, somewhat tendinous,
from the outer surface of the external condyle of the
femur, just below the external head of the gastrocnemius.
The fibres go to form a thin, flattened muscle, broadly
spindle-shaped, and about three centimetres long, with
its tendon merging with the spreading fascia of its outer
aspect. This tendon is rather wide for the most part
above, and very thin. It commences at the lower apex
of the muscle, and takes a straight course to the supero-
internal angle of the tibial cartilage, thus really passing
in an oblique line across the back of the leg. It be-
comes gradually smaller as it nears this cartilage, which
it passes through, superficially, to its outer side ; then
crossing to the tibio-tarsal joint, passes through a special
canal in the hypotarsus, in the median row to the inner
side. Its course is now directly down the back of the
tarso-metatarsus, under the annular ligament of the sole
of the foot, and a cartilaginous block beneath it to hold

Le Jiechisseur per/ore (pt.). Yicq d'Azyr, p. 283, No. 3.
Le muscle perforant et perfore (pt.) Cuvier, p. 558.
Zusammenziehei- der Zehen (pt.) Merrem, p. 161, No. 7. ^
Flexor perforatus digiti interni. Wiedemann, p. 102.

„ ,, „ „ Tiedemann, § 309.

Zweiter hinterer Kopf des durchbohrten und durchbohrenden Zehen-

beugers. Meckel, Archiv, p. 276, No. 9.
Erster hinterer Kopf des durchbohrten und durchbohrenden Zehen -

beugers. Meckel, System, p. 380, No. 1.
Flexor digitorum sublimis s. perforatus (pt.). Gurlt, p. 31.
Flexor perforatus s. longus dig. (pt.). Owen, Apteryx, p. 295.
Flechisseur perfore, couche superficielle (pt.). Gervais et Alix, p. 37.
Flexor digiti II. et III. {perforans et perforat\i,s) (pt.). Quenner-

stedt, p. 40 ; Neander, 24.
Flexor ])erforatus digitortmi (ausserer Kopf). De Man, p. 131,

No. 20.
Flexor perforans et perforatus digiti II. Gadow, No. 34.
Flexor perforatus et perforans digiti interni. Watson, p. 122."

Fjg. 64 his. — Pelvic limb, right side, oiGcococcyx californianus, same specimen as is
shown in Figs. 62 his and 63 ^^'.9. Still other muscles have now been
removed, and the amhiens is in full view, while other interesting parts
are brought into sight. Life-size, by the author, from his own
dissections.

232 THE MYOLOGY OF THE RAVEN.

these teiiclons in place, over the trochlea, to the middle
of the shaft of the second phalanx of the second toe.
Here it makes fast to the sides of this l3one as well as
forming a tul)nlar sheath for the passage of the deep
flexor.

When this mnscle alone contracts, hy r)nllLno' on the
second phalanx of the second toe it tends to flex it ;
when all the flexors of the limb act, it no doul)t is an
eflicient auxiliary to the same end.

117. The jiexor longus hallucis^ is a strong muscle
at the back of the leg, overlain l)y two other flexors of
the toes, presently to be described, which take origin
from its side. It arises l)y two very distinct tendinous
heads : the one, a median one from the posterior aspect
of the femur, immediately between the two condyles :
the other from the outer aspect of the external condyle
of the femur, just below the origin of the preceding
muscle. The deep flexor passes up between these two
heads, or rather passes down from its origin.

The Jiexo)' longus hallucis now forms a large fusiform
muscle at the back of the tibia overlying the deep flexor.
At the lower third of the leg it comes to a gradual point,
to terminate in a strong tendon, that passing beneath
the more superficial flexors, in a special canal on the
outer side of the tibial cartilage, goes through the
outer and large canal of the hypotarsus, next to the
shaft and directly down to the apex of the accessory

^ "53i. M, FLEXOR HALLUCIS LONGUS.

Flexor hallucis longus. Gurlt, p. 31.

De Man, p. 133, No. 22.
Garrod, F.Z.S., 1872, p. 363.
Flexor digitorum communis profundris (pt.). Quennerstedt, p. 45.
Le flcchisseur profond du pouce. Alix, p. 459.

Flexor perforans digitorum (pt.). Watson, p. 124." (Gadow, loc.
cit., p. 197.)

N

THE MUSCLES OF THE LOWER EXTREMITY. 233

metatarsal bone. It winds round to the inner side
of this, fits snugly in its groove beneath, then passes
along on tlie under side of the sliaft of the basal joint
of hallux, to l^ecome finally inserted into the tubercle
at the inferior proximal end of its ungual phalanx.

This muscle is a powerful flexor of tlie hind toe or
hallux. When it contracts, however, it also acts upon
other flexors of the toes, soon to be alluded to, so there
is not absolute independence of action here.^

^ Of the flexor ferforatus indicis secundus jjedis and the flexor
longus hfjdlucis, as they occur in Geococcyx californianus, I have said
that : —

" The flexor inrfo'vatus indicis secuyidus pedis (Fig. 63 his) is
even a better developed muscle than I found it to be among typical
Corvida;, some of which I have recently dissected, and it is fully
as well individualized.

" It arises from the fascia at the outer side of the knee-joint, and
from the contiguous surface of the external condyle of the femur.
Here it receives the anastomosing fibres of the extremity of the
tendon of the amhieyis.

" The muscle is fusiform in shape and accurately moulded on the
flexor it covers at its side. Its tendon in descending the leg is thin
and ribbon-like. At the ankle it passes through the tibial cartilage,
and crossing the joint goes through, with the second tier of tendons,
the cartilaginous cap on the back of the hypotarsus of the tarso-
metatarsus. Passing down behind this latter bone, and through the
annular ligament in the sole of the foot, it proceeds to the under side
of the second toe, beneath the second phalanx of which it expands to
form a tubular sheath for the passage of the deep flexor, while at
the same time it becomes attached to the side of this joint of the toe
in question.

" The carneous portion of this muscle in the leg is to the otiter side
of the loop for the hiceps flexor crucis, and, owing to the fact that it
varies in form and size in different birds, it is as well to bear this in
mind.

" The flexor longus hallucis (Fig. 64 bis) has two separate heads,
the one coming off from the outer side of the external condyle of the
femur, and the other, far more fleshy, arising from the posterior
aspect of the same bone between the condyles. Above, this muscle

234 THE MYOLOGY OF THE RAVEN.

Some ten years ago the late Professor Garrod con-
tributed to the Proceedings of the Zoological Society
a paper setting forth some of his very important and
interesting investigations upon the deep plantar tendons
in different birds.

He shows that there was considerable diversity among
these tendons in the class, both in their behaviour in the
sole of the foot, as well as at the back of the tarso-
metatarsus.

The tendons of the Jiexor perforans digitorum, and
the present one, sometimes cross each other in certain
birds in a peculiar though constant way ; at others,
these two tendons are united by a descending fibrous
vincidum. " In all cases this vinculum is alw\ays
directed downwards from the hallux muscle to the
digits muscle, so that, when the tendon of the jiexor
perforctns digitorum, alone is pulled upon, the three

is overlapped by the more superficial flexors, while in turn it has
beneath it the Jiexor perforans digitorum profundus. About half-
way down the leg it gives way to a strong tendon, which, passing
deep in the tibial cartilage, crosses the ankle-joint to pass through
the outer canal of the osseous portion of the hypotai-sus of the tarso-
metatarsus. Down the back of the shaft of this latter bone the
tendon exhibits a disposition to develop an osseous rod in its con-
tinuity, but this does not actually occur in my specimen. It lies in
this region just above the tendon of the deep flexor, and, immediately
above the sole, makes a fibrous connection with it of some extent.
This fibrous ' vinculum ' is in no way oblique as it is desci-ibed by
Garrod for many birds, but passes directly from one tendon to the
other for about eight millimetres, and were it not known that it as
a rule passes obliquelyy»"o»i the Jiexor longiis licdlucis, it would be
quite impossible here to designate which tendon was responsible for
the connection.

" In the foot the long tendon of the hallux passes in the usual
way to become inserted on the tubercle at the under side of the
proximal end of the ungual phalanx." (See 120 of Bibliography at
the end of this volume.)

THE MUSCLES OF THE LOWER EXTREMITY. 235

anterior digits alone are flexed ; but when the flexor
longus hallucis is put in action, the digits as well as
the hallux are simultaneously flexed " (Coll. Memoirs,
p. 291). No such vinculum exists in the Eaven, and
in this particular it agrees with the Passeres and
Upupa epops. Further studies in this direction will
be very acceptable.

A few years ago I published the following remarks in
reference to the variations to be seen in the arrano^ement
of these plantar tendons, and the uses that could be made
of them in avian taxonomy (124 of Bibliography). I
said that " both Professor C. J. Sundevall and Professor
Garrod have paid considerable attention to the dis-
position of these tendons in the feet of birds. If I
mistake not, the first-named author was the writer who
originally invited attention to the fact that the tendon
of the flexor longus hallucis was completely independent
of the tendon of the flexor perforans digitorum pjro-
fundus in the Passeres ; and in view of this fact he
grouped these birds together, and as the Hoopoe {Upupa)
exhibited the same condition, he included that form with
them. Garrod pushed the matter much further, however,
and made some very extensive dissections upon the deep
plantar tendons in a great many different orders of birds.

" Irrespective of the plan of the foot, in all birds, in so
far as its digits are concerned, there are two muscles
present in the leg, which, arising from the tibia and
fibula, send each a tendon to the toes a,^ flexors : these
muscles are the flexor longus hallucis and the flexor
perfoi^ans digitorum profundus. In passing through or
over the hypotarsus of the tarso-metatarsus, at the back
of the ankle-joint, the tendon of iho, flexor longus hal-
lucis is either superflcial or external to the tendon of the
other flexor mentioned.

^^^^"'a^.

J^Hu-Cu!^

Ci6-ltll C«y.7

Fig. 65. — Outer aspect of the left pelvic limb of a Raven, showing the fourth or
deepest layer of muscles of the thiglx and leg. The under side of the
obturator intcrnios may be seen through the " obturator space," and its
tendon surrounded by the gemellus attached to the trochanter of the
femur. The tibia has been rotated in order to bring its posterior
surface into view, and show the muscles there found; the "tibial
cartilage " is seen at its distal extremity. Life-size, by the author, from
his own dissections.

Fig. 65 bis. — Pelvic limb, right side, of Gcococcyx calif or nianus, same specimen
as is shown in Figs. 62 his, 63 his, and 64 his. The deepest muscles of
all only are left, and the tendon of insertion of the ohturator internus
is distinctly seen. Life-size, by the author, from his own dissections.

238 THE MYOLOGY OF THE RAVEN.

" This fact is useful to be borne in mind in identifying
these tendons in our dissections. Now after they pass a
short distance down the back of the tarso-metatarsal bone
their behaviour in different birds is quite diverse, and
a few examples of it will be here presented in order to
show that when our knowledge becomes more full in the
premises, the character wdll prove a useful one in classi-
fication of birds ; and also it is hoped to induce those
interested in the science of anatomy to undertake and
carefully record researches upon this subject. Before
arriving at the podal phalanges, and in the sole of the
foot of any bird, these tendons divide into a sufficient
number of slips to be distributed to the former, one slip
going to each toe. The method of division is the
same for the bulk of avian families, and the more uni-
versal type is well exemplified in the Common Chicken
(Gallus).

"Fig. 65 quat. of this paper shows this arrangement
in the Fowl, and there we observe that the tendon of the
flexor to the first toe is external to the tendon of the
flexor pc^florans digitorum 'profitndus as it passes the
ankle-joint.

" At the back of the tarsus it crosses the latter super-
ficially, and then passing directly to the under side of the
hind-toe it becomes inserted into the base of its ungual
phalanx.

" Now the larger tendon of the f. j)- digitorwn j)ro-
fundus, after arriving at the sole of the foot, trifurcates,
and a slij) is sent to the under side of each anterior toe,
where passing forwards they too become inserted at the
bases of the ungual digits of the respective phalanges.
Just above this trifurcation the tendons of these two
muscles however, are connected, and that by a fibrous
vinculum {¥\g. 65 quat., Vn) which passes between them.

THE MUSCLES OF THE LOWER EXTREMITY.

239

" The fibres of this vinculum come off from tlie tendon
of the jiexor longus halliicis, and pass downwards to
soon merge with the fibres of the tendon of tXia Jiexor
jDerforans digitorum profundus at the crossing.

Fig. 6s tei

Fig. 65 quat.

Fig. 6^ quin.

Fig. 65 ter. — Right foot of Mcgalcevii asiatica, showing an arrangement of the

plantar tendons.
Fig. 65 quat.—^i^h foot of Gallus bankiva. Vn, vinculum running downwards

from the outer hallucial tendon to the inner digital common tendon.
Fig. 65 qioin. — Right foot of Apteryx mantelli. (All three figures copied by the

author from Garrod. )

" The strength and size of the vinculum is different
in nearly every group of birds where it is present. In
Fig. 65 his, at a, I have shown the position and size of

240 THE MYOLOGY OF THE RAVEN.

this vinculum as I found it to be present in the Ground
Cuckoo.

" In order to show how different this disposition of these
plantar tendons may be, I quote Garrod's description of
his dissection of the foot of a specimen of Megalcsma
asiatica (Fig. 65 ter.). He says, in this bird ' the two
tendons descend behind the ankle as usual, having their
origins typical. There is nothing peculiar till they have
descended two-thirds down the tarso-metatarse. About
opposite the middle of that bone the fiexor longus
liallucis sends a vinculum downwards as in the Fowl, to
join the tendon of the, flexor i^Gvf or cms digitorwn. Just
above the metatarso-phalangeal articulation the tendons
become arranged for distribution in a most uncommon
manner. The tendon of the flexor iK'rforcms digitorum
does not S2:)lit up, but runs to one digit only — namely,
the third toe, which is the outer of the two that are
directed forward. It is covered superficially by the
flexor 2^erforans digitorum, just as that latter muscle is
splitting up to be distributed to the hallux as well as to
digits 2 and 4. In these birds we have, therefore, the
flexor longus liallucis arising from the lower, surface
of the femur only, running through the ankle at
the outer side of the other deep tendon, and sending
a vinculum downwards — all of which are special cha-
racters of that muscle only, it being distributed to
three toes, whilst the flexor perforaiis digitorum only
supplies one.'

" In Fig. 65 quin. the arrangement in the Ajyferyx is
well shown, where the vinculum is so large as compared
with this band in the Fowl, that it almost appears upon
sight that the two fiexor tendons are fused into one
common one, at the point of union.

"A close study of this arrangement, however, will throw

THE MUSCLES OF THE LOWER EXTREMITY. 241

much light upon the variations as they are to be found
in the less tyjDical birds in this respect."

118. The Jiexor perforatus annularis primus pedis ^
arises from the iniier side of the belly of the Jiexor
longus hallucis, about a centimetre below its median
head, some of the fibres of its flat tendon being un-
doubtedly carried up to the insertion of the latter
muscle. It passes directly down the back of the leg
as a long, laterally compressed, rather slender muscle.

^ In introducing the term " annularis " into the name of this
muscle, the writer does so simply with the view of impressing upon
the mind the relation or position the toe bears with reference to the
corresponding finger of the hand, i.e. in five-fingered and five-toed
vertebrates. A distinctive name is pretty sure to do this. I am
greatly pleased with the names Gadow has conceived for these flexor
muscles in the feet of birds, and if what the names signify holds true
for the class, they are deserving of veiy general adoption. No one
will question the boon it would prove to be to have a commonly
accepted nomenclature for them (No. 118 is 51c of Gadow).

"51c. M. FLEXOR PERFORATUS DIGITI IV.

Le JlecliissetiT commun {on ferfore). Vicq d'Azyr, 283, No. 3 ;

Cuvier, 558.
Flexor jjerforatus digili externi. Wiedemaun, p. 104.
,, ,, „ „ Tiedemann, § 311.

„ „ ,, ,, Watson, p. 121.

Langer Zehenbeuger ; innerer Kopf. Meckel, Archiv, 277, No. 4.
Einer der Spulmuskeln. Meckel, Syslem, 382-386, No. 2.
Flexor eonwiunis quatluor digitorura s. subliniis (pt.). d' Alton, p.

37.
Flexor 2)er/oratus (Tpt.). Reid, p. 144.
Flexor suhlimis s. i^erforatus digiti quarti. Gurlt, p. 31.
Flexor perforatus of the outer toe, s. longus digitorum (pt.). Owen.
Flexor perforatus digitorum (ausserer Kopf). De Man, 131, No. 20.
Flechisseur peifore, coucheprofonde (pt.). Gervais et Alix, p. 36.
Flechisseur superficiel du doigt externe (ou quatrieme doigt). Alix,

pp. 458 et 460.
Flexor digitorum suhlimis. Quennerstedt, p. 42 ; Neander, p. 25.
Flexor perforatiis digiti IV. Gadow, No. 38."

E

242 THE MYOLOGY OF THE RAVEN.

It terminates in a flattened tendon at the lower third
of the tibia, which is closely applied to a similar
tendon coming from the flexor j)erforatus medius
jwimus pedis, and the two pass together through the
tibial cartilage, rather to its outer side, in a special
canal. Crossing the ankle-joint, they also pass together
through a canal in the hypotarsus situated at its upper
and outer angle.

At the back of the shaft of the tarso-metatarsus
these tendons part company, the tendon of the flexor
perforatus medius secundus jjiet/i's passing Ijetween
them obli<]uely.

The tendon of the muscle now under consideration
makes straio-lit for the under side of the outer toe.
Here it forms a tubular sheath, through which the deep
flexor runs, and opposite the short shafts of the l^asal
phalanx and the joint next beyond, it sends down
on either side a tendinous slip, which in each case
attaches to these bones. Thus fulfilling, we see, the
part, too, of a secundus flexor of this toe.

119. The flexor 'perforatus medius prm?/s pedis ^
arises from the inner side of the carneous portion of

1 Referring to the synonymy of this muscle as compiled by Gadow,
we are enabled to record the following upon the subject [loc. cit., p.
188): —

"51b. M. FLEXOR PERFORATUS DIGITI III.

Le Jlechisseur conmiun (ou 2^erfore) (pt.). Vicq d'Azyr, p. 283, No. 3 ;

Cuvier, p. 558.
Zusammenzieher der Zehen (pt.). Merrem, p. 161, No. 7.
Flexor j)erforatus digiti medii. Wiedemann, p. 103.
„ „ ,, „ Tiedemann, § 310.

„ „ ,, „ Watson, p. 121.

Langer Zehenbeuger, ausserer Kopf. Meckel, Archiv, p. 277,

No. 4.
Einer der Spulmuskeln. Meckel, System, pp. 382-386, No. 2.

THE MUSCLES OF THE LOWER EXTREMITY. 243

the last-described muscle close up to its head. It is
considerably smaller in size, and its outer fascia sooner
terminates in its flattened tendon. This takes its course,
as I have already described, closely applied to the
jiexoT 'perforatus annularis primus pedis, through
the tibial cartilage and hypotarsus of the tarso-meta-
tarsus. Arriving at the palmar aspect of the basal
joint of the third toe, it sends down a slip to either
side of its shaft, which thus forms a guide for the
passage of the two deeper flexors which glide between
them.

The carneous portions of the three last-described
muscles are quite intimately bound together by a
close-fitting and firm fascia. So that there is con-
siderable unison in their action, but, as we have seen,
the course of their tendons is more or less independent.
If it were possible to keep the distribution and in-
sertion of these tendons in our mind, it would not be
far out of place to speak of their common muscular
portion as the jlexor 2^e7]fo7^atus pnnnius jyedis.

120. The Jlexor j^e^Joratus medius secundis ^9ec?{.v ^

Flexor communis quattuor digitorum s. suhlimis (pt.). d' Alton, p. 37.

Flexor digiti secundi et tertii brevior (pt.). Gurlt, p. 31.

Flexor perforatus s. longus digit07-um i^t.). Owen,

Flexor perforatus digitorum (innerer Kopf). De Man, p. 131, No.

20.
Flechisseur perfore, couche p/rofonde (pt.). Gei"vais et Alix, p. 36.
Flechisseur de la deuxieme p)halange du doigt interne (Struthio) ; du

doigt troisieme (Ardea). Alix, pp. 457 et 460.
Flexor digitorum suhlimis (pt.). Quennerstedt, p. 42 ; Neander

p. 25.
Flexor perforatus digiti III. Gadow, No. 37."

^ The synonymy of Gadow is thus recorded {loc. cif., p. 192): —

** 526. M. FLEXOR PERFORANS ET PERFORATUS DIGITI III.

M. sextus circa tibiam et fibulam. Steno.

Flechisseur perforant et p)erfore (pt.). Yicq d'Azyr, 284, No. 4.

R 2

244 THE MYOLOGY OF THE RAVEN.

is one of tlie largest of these perforated flexors the
tendons of which go to the toes.

It is found at the antero-external aspect of the
leg, and arises by a strong fascia that merges with
the fascia of the knee-joint. It also arises by a strong
tendon, in common with the tendon of the flexor
perforatus indicis secundus j^edis, from the outer side
of the external condyle of the femur. The muscles
of the anterior aspect of the til^ia also blend with it,
more or less above, by a common fascia ; while below
we find it has a semi-tendinous attachment with the
lower end of the filnila, and a line on the tibial shaft
for a short distance below it.

Le muscle perforant et perfore (pt.). Cuvier, p. 553.

Fingerschliesser. Merrem, p. 161, No. 8.

Flexor ferforans et perforatus digiti medii. Wiedemann, p. 102 ;

Tiedemaun, § 314; Watson, p. 123.
Zweiter vorderer Kopf des durchbohrten Beugers. Meckel, System,

p. 380, No. 1.
Erster vorderer Kopf des durchbohrten Beugers. Meckel, Archiv,

p. 276, No. 3.
Flexor digitorum suhlimis s. perforatus (pt.). Gurlt, p. 31.
Flexor perforatus {^t.). Reid, p. 144.
Flexor p)erforatus s. longus digitorum (pt.). Owen.
Der besondere Beuger des ersten Gliedes der dritten Zehe. Carus,

ErlauteruDgstafeln.
Flexor digiti II. et III. [perforans et perforatus) (pt.). Quennerstedt,

p. 40 ; Neander, p. 24.
Flechisseur perfore, couche superficielle (pt.). Gervais et Alix, p. 37.
Flechisseur de la troisihne pilialange du doigt interne (Struthio)

troisieme. Alix, pp. 457 et 460.
Flexor perforans et perforatus digiti medii. De Man, p. 134, No. 23.
M. flexor perf or ans et perforatus digiti HI Gadow, No. 35.
Flexor p)^iforans et perforatxhs digiti medii. Watson, p. 123."

Note. — The accounts of the flexors and extensors of the toes in
birds as given by Prof. Gadow in Bronn's Thier-Reichs can very
profitably be comj)ared with the musculature of the Haven,

THE MUSCLES OF THE LOWER EXTREMITY. 245

From this extensive origin, this hirge and fusiform
muscle, covering the fibular region, terminates at the
lower third of the leg in a strong and somewhat
flattened tendon. This passes obliquely and very super-
ficially through the tibial cartilage from without
inwards to the hypotarsus, where it passes through the
interno-posterior canal, then accompanies the other
tendons at the back of the metatarsus, to deviate and go
to the second joint of the middle toe, having perforated
the tendon of the basal phalanx en jKissant. Its method
of attachment to this prebasal joint is precisely the
same as for the corresponding tendons of the other
podal digits described above.

121. The flexor j^^^'^lforatus indicis jjrimus 2)<^dis^

1 This is the Jlexor per/oralus digiti II. of Prof. Gadow's nomen-
clature, and he gives the following synonymy for it (foe. e?7., p. 187): —

"51a. M. FLEXOR PERFORATUS DIGITI II.

Le Jlechisseur du doigt interne. Vicq d'Azyr, p. 289, No. 3.
Flechisseur commun (ou perfore) (pt.). Yicq d'Azyr (1) ; Cuvier,

558.
Flexor profundus phalangis I. digiti interni. Wiedemann, p. 105.

,, ,, ,, ., ,, „ Tiedemann, § 313.

„ „ „ ., „ De Man, p. 133, No.

21.
Einer der Spulmuskeln (1). Meckel, System, pp. 382-386, No. 2.
Flexor conmiunis quattuor digitoruni s. suhlimis (pt.). d' Alton, p. 37.
Flexor digiti secundi et tertii brevior (pt.). Gurlt, p. 31.
Flexor perforatus (pt.). Reid, p. 144.
Flexor perforatus s. lotigus digitorum (pt.). Owen.
Flechisseur per/ore, couche p/rofonde (pt.). Gervais et Alix, p. 36.
Flechisseur de la deuxihne phalange dudeuxihne doigt. Alix, p. 460.
Flexor digitorum suhlimis. Quennerstedt, p. 42.

,, ,, ,, Neander, p. 25.

M. perforatus digiti II. Gadow, No. 36.
Flexor pjerforatus digiti interni. Watson, p. 120."

Gadow seems to have set to work at his analysis of the muscula-

246 THE MYOLOGY OF THE RAVEN.

is situated posterior to the preceding, and is a long,
slender, somewhat flattened, fusiform muscle, whose
office it is to assist in flexing the second toe. Laterally
its fascia blends, more or less, with the flexors on either
side of it ; still, the muscle is well-individualized, and
fully entitled to a separate description and name. It
arises by a thin and rather broad tendon, in common
with the flexor 2^^^fl'^'*^<^^^^'^ medius secundus ^9e(iiA',
from the external condyle of the femur. Below, it
terminates in a strong though slender tendon, before
the last-named muscle does, and passes through the
tibial cartilage and hypotarsus of the metatarsus in
precisely the same manner, though in each case in a
canal deep to it.

In accompanying the other flexor tendons down the
back of the shaft of the tarso-metatarsus, it is found
rather to the inner side. Within the region of the
sole of the foot it is situated very deep, and just within
the accessory metatarsal bone. It finally becomes
attached to the sides of the basal joint of the second
toe, in the same manner as the other perforated tendons
in the other toes, the secundus and deep ones"passing
throuorh its median bifurcation.

122. The flexor j^Grforans digitorum 2^^'ofu7idiis^ is,
in point of situation, the deepest of all the flexor

ture of the feet in Aves very much in the same manner as did the
present writer, and his vahiable chapter on the subject in the work
we have so often qvioted in the present volume is well worthy of the
closest study and perusal.

^ We have here a very important muscle of the leg, and one that
was thoroughly investigated by Garrod at the time he was in search
of structviral characters among birds that could efficiently be em-
ployed as a means in classification.

Gadow has collected together the following synonymy for it, and
in his work in Bronn's Thier-Eeichs gives quite a full resume of the

THE MUSCLES OF THE LOWER EXTREMITY. 247

muscles at tlie back of the tibia (see Figs. 46, 52, 59,
60, and others).

Its highest origin is by a strong, fleshy head from
the posterior aspect of the external condyle of the
femur. Two other fairly defined heads of this muscle
come off from the tibia behind, just below its summit ;
the inner one being close to the origin of the soleus.
This deep flexor also arises from the major portion of
the posterior surface of the tibial shaft (Fig, 52) as
w^ell as from the fibula. In form, it is flattened, with
a strong fascia covering its exposed or external surface,
extending nearly its whole length from the femoral
head. At the lower third of the tibia it terminates

variatioQS the ^flexor dig. jwofundus takes on in the various groups
of birds (Joe. cit., pp. 193-196) :—

"53a. M. FLEXOR PROFUNDUS S. PERFORANS.

M. secundus j)osterior digitos movens. Aldrovandi.

M. octavus 2Josterior circa tibiam et fihulcmi. Steno.

Le Jlechisseur 2^6'Kfo'>'(i'fi't ■ Vicq d'Azyr, p. 284; Cuvier, p. 558.

Zweikopfiger Muskel. Merrem, p. 161.

Flexor profundus s. perforans trijidus. Wiedemann, p. 104.

Flexor profundus s. perforus trifidus. Tiedemann, § 312.

Durchbohrender oder tiefer durchbohrender Beuger. Meckel, System,

p. 386-390, No. 3 ; Archiv, 278-279, No. 5.
Flexor profmdus s. perforans ; flexor communis IV. dig. s. suhlimis

pars post, d' Alton, p. 38.
Flexor digitorum longus s. perforans. Gurlt, p. 31.
Flexor perforans digitorum. Owen, Cyclop., p. 297 ; Aptcryx, p. 295 ;

De Man, p. 130, No. 19 ; Watson, p. 124.
Flexor digitorum communis profundus. Quennerstedt, p. 45 ;

Neander, p. 27.
Flexor perforans. Reid, p. 144.
Flechisseur profond. Gervais et Alix, p. 35.
Flechisseur pyrofond, perforant ou commun des trois doigts j)ro])re7nent

dits. Alix, p. 459.
Flexor irrofundus. Garrod, Proc. Zool. Soc, 1872, p. 363 ; Gadow,

No. 39."

248 THE MYOLOGY OF THE EAVEN.

in a powerful and flattened tendon, which passes,
nearly in the median line, through the tibial cartilage,
deep to all the other flexors. In traversing the hypo-
tarsal prominence it selects the large outer canal next
to the shaft; and in emerging from it, its tendon, in
passing down the back of the shaft of the tarso-meta-
tarsus, is nearest the bone. It also develops in it, in
this locality, a strong osseous rod, just below which
its trifurcation takes place. Each division as it leaves
the main trunk in the sole of the foot, proceeds directly
to one of the anterior toes, to pass its entire length
along its palmar aspect. In eff"ccting this, it lies close
to the several joints, and perforates the other flexor
tendons in a manner already described. Arriving at the
ungual phalanges, in each case, the tendon attaches
itself to the proximal tubercle at the l)ase upon its
under side. These several tendons are also held in
position by strong fasciae which bind them to the under
side of the toes ; and these must be slit up with our
scalpel before they are exposed.

This muscle, as may be seen both from its origin
and insertion, is the most powerful of all the " flexors
of the toes ; the others being, more correctly sj^eaking,
simply auxiliaries to its action.

After the integuments have been removed, the form
of the leg is quite conical, being, however, somewhat
compressed laterally, while its base may be said to be
at the knee-joint, and its apex at the condyles of the
tibia.

All the flexors we have described above are more
or less intimately connected together by binding fascia,
and this applies also, to a great extent, to the group
of muscles upon the anterior aspect of this division of
the pelvic limb ; while the whole seems to be enveloj)ed

THE MUSCLES OF THE LOWER EXTREMITY. 249

in a firm slieatli of connective tissue, which masks at
first sight the individuality of the several muscles.

For a centimetre or more above the tibial condyles
we observe their numerous tendons, as they start from
the lower apices of these different muscles ; while in the
division of the limb next l^elow, we find the flexors at
the posterior aspect of the tarso-metatarsal shaft snugly
wrapped in fascia, and bound to the bone as a sub-
cylindrical bundle of tendons, the several factors of
which enjoy perfect freedom of action in the longitudinal
direction. The extensor tendons and the short extensor
muscle and other elements are somewhat differently
circumstanced in front in the very nature of their re-
quirements, but here, too, a fascia surrounds them in
a similar fashion, drawing them up snugly by the shaft
of the bone.^

^ In the second volume of his Anatomy of Vertebrates, Professor
Owen describes for the Ajoteryx (p. 104) a, popliteus muscle, and says
it " is brought into view when the superficial muscles of the leg
which are inserted into the foot are removed. Its carneous fibres
extend from the fibula inward and downward to the tibia. It is of
relatively smaller extent than in the Cassowary." Professor Mivart
also alludes to the occurrence of this muscle in birds in his Elementary
Anatomy, but I must confess that after careful search in a number
of individuals, 1 have signally failed to find the popliteus in the
Raven. And I am rather inclined to believe that it will be found
to be absent in the higher groups of birds. It is present in the
Echidna.

Since writing the foregoing paragraph of this footnote, I find
the following remarks and synonymy in Gadow on the popliteus
muscle in birds (loc. cit., pp. 176, 177). He does not mention it as
occurring in the Corvidm.

" 44. M. POPLITEUS.

Le muscle po'plite. Yicq d'Azyr, p. 514.
Foplitceus. Wiedemann, p. 99.
Popliteiis. Tiedemann, § 303.

250 THE MYOLOGY OF THE HAVEN.

We find a well-developed popUteus muscle in Geo-
coccyx calif ornianus, where it is seen to arise from an
oblique line on the back of the tibial shaft below the
head of the bone, and the fibres converging to pass
upwards and outwards are inserted by a short tendon
into the corresponding aspect of the head of the fibula,
close to the superior fibres of insertion of the jiexor
perforans digitorum j^^'oftmdus.

The several flexor muscles of the pelvic limb of the
Eaven which we have been describing in the last few pages
can be compared with great profit with the corresponding
muscles as they occur in a bird with zygodactyle feet.
The species to which we have been referring for that
purpose in this work is Geococcyx calif ornianus, and, as
has already been noted, the writer presented some ob-
servations upon its myology in the Proceedings of the
Zoological Society of London for 188G. From that
paper I here republish my remarks upon these flexor
muscles as they were found to exist by me in Geococcyx,
in order that the comparisons can be made which have
been suggested above. 1 remarked that in Geococcyx

Popliteus. Owen,

„ Quennerstedt, p. 36.

,, Neander, p. 21.

Gadow, No. 28.
„ Watson, p. 116.

Kniekehlenmuskel. Meckel, Siji^tem, p. 369, No. 2 ; Archiv, p. 272,

No. 11.
Le pojilite. Gervais et Alix, p. 33.

" Ausserordentlich klein ist der M. jmpliteus bei den kletternden
Vogeln, auch bei Guculus, wahrseheinlich ganz feldeud bei Psittacus,
Picus, und Cypselus. Dagegen ist er recht anselmlich bei Capri-
mulgus, Strix, Buteo, Gallus, Numenius. Schvvach bei Anser, Grus,
Fulica, Larus, Gohjmhus, Ciconia, Serpentarius."

THE MUSCLES OF THE LOWER EXTKEMITY, 251

" the Jic'XOT 'pa'yfoTatus arMiuIcvris jwimus pedis (com-
pare Figs. 63 his with 65 his) in this bird seems to have
acquired a more central position on the back of the leg
than in many others that I have dissected, and, more-
over, its tendon, as will be seen from the figures, is quite
suj)crficial.

" It arises from between the condyles of the femur by
•a slight semitendinous slip, and from the side of the
Jiexor longus hallucis. The muscle itself is somewhat
of a fusiform outline and rather flat ; it lies to the iymer
side of the loop for the hicej^s. About one-third of the
way down the back of the leg it terminates in a small
though very long tendon, which, passing quite super-
ficially through the tibial cartilage and over the ankle-
joint and hypotarsus, runs in common with the other
flexors down the back of the tarso-metatarsus, turns to
the rear in the sole of the foot to become inserted on the
under side of the distal end of the basal phalanx of the
reversed digit. In the specimen before me this insertion
is to the outer side of the deep flexor, and the tendon is
not slit for its passage. Nothing could be more engaging
than the examination of these reversed tendons in the
sole of the foot of this Ground Cuckoo, for the greatest
nicety in accommodation has been accomplished as they
have gradually come to assume their present position.
No doubt some of the departures observed from the
more common arrangement of them are due to the
reversion of the digit in question.

" Strong, fibrous bands are so disposed in this jDlantar
region as to admirably hold the several groups of flexor
tendons in place, and at the same time they act as pulleys
for their guidance and afford correct application of the
force intended to flex the toes.

" We also have in Geococcyx an unusually large j/?ea?07^

252 THE MYOLOGY OF THE RAVEN.

'perforatus m,edins 'prmins pedi'^, wliich here arises by two
slips, an outer tendinous one, from the external condyle
of the femur, which has a common origin with other
muscles there arising, and is intimately connected with
the dense fascia about the front of the knee-joint ; while
the second slip arises from between the femoral condyles,
in common with other flexors that come off from that
point. The two heads are quite independent, but merge
with each other before they terminate in their common
tendon at the lower third of the tibial shaft.

" It passes through the tibial cartilage, overlain by,
but in close company with, the far more diminutive
and narrower tendon of the Jiexor perfo)-atus annularis
2)rhmis i^edis.

" When it arrives under the basal phalanx of the
median toe, the outer one of the anterior pair, it l)ifur-
cates to allow the other two flexor tendons to pass, while
the slips thus formed become attached to the sides of the
shaft of this j oint close to its distal head.

" It will at once be seen that Geococcijx californianus,
in common with the vast majority of birds, has no special
tendon devoted to the flexing of the second or prebasal
phalanx of the fourth toe (here the reversed one).
Provision is made for this in various ways in diff'erent
birds. Here, in the subject before us, a special slip is
thrown oft' for attachment from the deep flexor tendon
as it passes over the prebasal joint in question, which
slip virtually fulfils the function of a Jiexor perforatus
annularis secundus pedis, did such a muscle with an
independent tendon exist.

"The Jiexor p>erforatus medius secundus pedis, as in
all of the birds I have examined, is one of the best
developed perforated tendons at the back of the leg. Its
fascia of origin merges with the enveloping fascia about

THE MUSCLES OF THE LOWER EXTREMITY. 253

the knee-joint, while it also arises hy a strong tendon,
common to it and the Jiexor perforatus indicis secundus
pedis, from the external aspect of the outer condyle of
the femur.

" Finally, it is attached more or less by carneous
fibres down the shafts of the leg-bones to a point
below their middle, or rather the middle of the leg.
Below this the muscle terminates in a strong tendon,
which, taking an oblique course through the tibial
cartilage, passes as usual over the ankle-joint, through
the hypotarsus, and down the back of the tarso-
metatarsus.

" In the foot it perforates the more superficial flexor
of the basal phalanx of the outermost of the two anterior
toes, then in turn bifurcates over the prebasal joint to
allow the deep flexor to pass through, these bifurcations
becoming the insertions of this muscle, and they are
attached to the sides of the shaft of the second joint of
the toe alluded to, or the external one of the two in
front.

"As its name indicates, our next muscle, the flexor
pei'foratus indicis p>rimus p)edis, has its tendon at-
tached to the nether side of the basal phalanx of
the index digit, and consequently aids in bending
that toe.

" Above, as a flat, long muscle, it comes off by a thin
tendon from the external surface of the outer femoral
condyle, arising with the flexor p>erforatus medius
secundus p>edis.

"The flexor perforans digitorum profundus. This
muscle is deep to all the flexors, and is situated directly
on the posterior aspect of the tibia and fibula. It arises
by two heads, one from the upper part of the tibia im-
mediately below the overhanging rim of its summit.

254 THE MYOLOGY OF THE RAVEN.

and the other, smaller, from the back of the head of the
fibula. In the Corvidae there is a well-developed third
head, which comes off from above the fibular notch of the
outer condyle of the femur, and in these birds, too, there is
no fibular head to this muscle, but two tibial ones instead.
Geococcyx agrees, however, with most birds in having
this muscle attached nearly the whole length of the
posterior aspect of the shaft of the tibia by lightly
attached carneous fibres.

" About a centimetre above the tibial cartilao^e it
terminates in a strong subcompressed tendon, which,
passing beneath the cartilage referred to, crosses the
ankle-joint in front of all the other fiexor tendons, and
then passes through the inner of the two longitudinal
perforations of the hypotarsal apophysis of the tarso-
metatarsus. Down the posterior aspect of the shaft of
this bone the tendon still maintains its anterior position
and exhibits a predisposition to ossify. But this does not
actually take place in the specimen before me. Above
the distal trochleso, it makes the fibrous connection with
the tendon of the fiexor longns haUueis already described.
Once within the limits of the post-trochlear space, the ten-
don of this muscle behaves in a manner common to most
birds — that is, it quadrifurcates, and each branch takes
a course close up to the joints on their plantar aspects,
and running through the slits in the perforated tendons
pass in each case to the ends of the toes, where they
become attached or inserted upon the infero -proximal
tubercles of the ungual phalanges. The one passing
along under the fourth digit sends up a slip which is
attached to the under side of the shaft of the prebasal
joint, thus making good the deficiency here for what
is represented in the other toes by an independent
tendon.

THE MUSCLES OF THE LOWER EXTREMITY. 255

" The tibial head of this muscle at its origin is
directly covered by the soleus, while in the aperture
existing between the two heads w^e can discern the
j^opliteus.

" These flexor muscles, and others on both the front
and rear of this limb, are moulded upon each other in a
manner that can only be justly appreciated by a personal
examination. In some the connections are quite feeble,
the intervening tissue being easily separable, while in
others the intimacy is very close, and great care is de-
manded on the part of the investigator to see that the
separation is made along the proper divisions.

" I am convinced from my studies that a greater
difference is to be found among the various muscles of
birds than we have ever accredited them with, and this
fact leads me to believe that the day will come when
these differences can be called into play in taxonomy with
excellent effect. Perhaps if the myology of the leg is
examined as carefully as Professor Garrod examined the
muscles of the thigh in this class, fully as many interest-
ing and valuable distinctions will come to light.

"As we pass the muscles we have described for this
limb of Geocoecyx in review, it will at once be recognized
that the list is unusually complete. All the ordinary
muscles of the thigh are present as found in birds, and all
highly developed. In the leg marked specialization and
organization are everywhere evident, while exceptional
muscles are here, too, fully represented.

" This complexity by no means diminishes as we
proceed towards the foot, for the arrangement of the
tendons as they course down the tarso-metatarsus and
the special musculature of this division of the limb is
manifestly indicative of high organization.

" Finally, we have the complex insertional extremities

256 THE MYOLOGY OF THE RAVEN.

of the intricate system above laid before us in the foot ;
and the most exquisite examples of adaptation, compact-
ness, and final requirements are to be seen throughout
the structure on every hand.

" The Jiexo?^ hrevis hallucis is an exceedingly interest-
ing little muscle. It arises from the side and the lower
margin of the inner aspect of the hyj^otarsus of the
tarso-metatarsus, and from some of the shaft of this
bone immediately below. The fibres converge to termi-
nate in a small tendon, which, passing down the postero-
internal aspect of the shaft, goes to the inner side of the
basal joint of the hallux, about which it winds, to finally
become inserted on its under side, at the proximal ex-
tremity of this joint, just a little beyond its articulation
with the tarso-metatarsal troclilea.

" Thus it will be seen that this little muscle is entirely
devoted to assist in flexing the hallux. Its mesial fibres
meet those of the muscle next to be described, down the
mid-longitudinal line of the shaft of the bone which
gives it origin.

" Equally engaging with the last is another still smaller
muscle, the extensor hrevis annulcuns, on the opposite
side of the same bone. Here we find its origin is' much
the same as the flexor hrevis hallucis, coming off from
the external aspect of the hypotarsus and the shaft below.
It soon terminates in an extremely delicate little tendon,
which, passing directly down to the fourth or reversed
toe, becomes inserted on the supero-inner aspect of the
basal phalanx of this digit.

" By its contraction it will act as a direct extensor of
this toe, a requirement no doubt made necessary through
the feeble manner in which this digit is now served by
the slip which goes to it from the common extensor of
these phalanges.

THE MUSCLES OF THE LOWER EXTREMITY. 257

" This tendon of tlie short extensoi^ gets its leverage
by the fascia which circuhxrly binds down all the tendons
of the flexors and extensors, just above the sole on the
one side, and which passes above the distal trochlear on
the other."

IX. THE MUSCULATURE OF THE TRUNK.

A PECTOKAL and a pelvic limb may now be removed
from tlie same, say the left, side, and tlie opposite limbs
are allowed to remain, in order that we may compare
the relations existing 1;)etween their mnscles and the
muscles of the trunk, which are to be descrilied in the
present section. Next, we carefully remove the entire
remaining integuments from the cadaver of our specimen,
cutting the dermal muscles as w^e do so, close to their
origins. In preparing the neck for our investigations, we
should carefully remove the trachea ; all superfluous fat
and connective tissue ; and the vessels and oesophagus.
At its cephalic extremity, we clear away the entire
hyoidean apparatus, and with it any other structures
that may stand in the way of our complete view of the
method of attachment of the neck-muscles to the base
of the cranium.

In speaking of the muscles of the vertebrae. Sir
Richard Owen says that " the muscles of the cervical
region are the most developed, as might be expected
from the size and mobility of this j^art of the spine ;
the muscles which are situated on the dorsal and
lumbar regions are, on the other hand, very indistinct,
feeble, and but slightly carneous ; they are not, however,

THE MUSCULATURE OF THE TRUNK. 259

entirely wanting. In the Struthious and short-winged
sea birds, in which the dorsal vertebrae are unfettered by
anchylosis, these muscles are more fleshy and distinct,
most so in the Ai^teryx'' [Anat. of Verts., vol. ii.
pp. 84, 85).

From the fact that none of the muscles in the back
of such a form as the Raven exactly correspond with
the dorsal muscles of the Mammalia, and certainly not
with the highly differentiated groups, or the several
layers, as we find them in man, it is more than prob-
able that in the present instance I shall be called uj)on
to curtail and simplify their nomenclature, and bestow
upon certain ones provisional names, which they may
wear until such times as our knowledge of the myology
of the Yertebrata in general is far more extensive than
it is at the present time.

AVe may hope for some light upon the question of
such homologies only through exhaustive and comparative
studies of the entire muscular systems of the Struthious
forms of existing birds, and such other types as Echidna,
the Duckbill, certain Fish forms, with such living Reptiles
as are known to approach these in their structure. At
the best, the unravelling of these homologies is an ex-
ceedingly difficult task, and one that demands all our
patience, as well as our keenest insight into morphological
problems. Not a little has already been accomplished in
the direction I point out, but very little as compared
with the enormous amount of exhaustive comparative
work that still remains in such fields, for the hands
and minds of the generations to come.

We find in most existing Chelonians that these mus-
cles of the dorsum have almost entirely disappeared ;
while among the Snakes they are highly developed both
as regards complexity and differentiation. In living

s 2

de^i'/no'iluuuig

Scvcro - Ztcnv-

ijilut'cus mirturULS

•ye^mitcmlcnosas.

^l^^fatov C^cizJLcLoe^.

Fig. 66. — Dorsal aspect of the skeleton of the trunk in a Raven. Designed to
show the origin and insertion of certain muscles thereto pertaining.
Drawn life-size, by the author, from his own dissections.

THE MUSCULATURE OF THE TRUNK. 261

Teleosts among fishes they form a bulky mass that goes
to be inserted at the l)ack of the cranium.

No less interesting is the study of the group of
caudal muscles in this Raven, as compared with several
of the forms suggested above. In a bird as high in
the scale of organization as our present sul)ject, of
course, these muscles now all subserve the special
movements of no less important an instrument to
the performance of flight than the tail. With many of
our existing Lizards, however, the tail must be consi-
dered more in the lio;lit of an ornament rather than an
essential part in the economy. For instance, our little
Anolis seems to get along quite as well when this
appendage is broken off short with his body as he
does with it ; whereas I dare say such a curtailment
in the ArchcBoptc/i-yx of the Jurassic would have proved
a very serious accident, if not a fatal one in many
cases.

Again, even in the Mammalia, as in the ring-tailed
Monkeys, we find the tail a very useful and important
organ, and in them the caudal muscles and their ten-
dons are powerfully developed to meet the special ends
of the act of caudal prehension.

Splendid demonstrations of this group of muscles in
existing Lizards are to be seen in Mr. Mivart's memoir
"On the Myology of Chaimdeon parsohii'' {F.Z.S.,
December 6, 1870, p. 850), and his "Myology of
Iguana tuber culata" {ibid., 1867, p. 1QQ); as well as
in Mr. Alfred Sanders's " Myology of Liolepis belli "
(P.Z.S., 1872, p. 154), and the " Myology of Phryno-
soma coronatum," by the same author {P.Z.S., 1874,
p. 71).

In speaking of these muscles in a general way, Mivart
says that " A whole series of muscles may be developed

262 THE MYOLOGY OF THE RAVEN.

which in man are entirely absent. These muscles are
the numerous muscles which move the tail, and which
may attain a vast bulk, as in the Cetacea and in Fishes.
To describe these muscles in detail would rather
come within the scope of a treatise on the compara-
tive anatomy of animals than within that of the
present work. Here, however, it may be stated that
the enormous coccyx of the Porpoise is provided not
only with the dorsal muscles which continue on backwards
the erector spinas (with its main divisions) from the
occiput to the tail end, but also possesses a ventral
muscular mass (extending forwards as far as the middle
of the thorax), which mass is divisible from above down-
wards into two antero-posteriorly extended masses —
together constituting, as it were, a ventral (and here
sub vertebral) reflection of .the erector spinse. The same
appearance occurs in some Reptiles and in Tailed
Batrachians, where the ventral muscles of the tail repeat
below, the dorsal masses above. But these Batrachian
caudal muscles are not subverteljral — not the con-
tinuation backwards of subvertebral ones of the trunk,
but direct continuations backwards of the abdominal
muscles, as is also the case in most Fishes " {Elem. Anat.,
pp. 323, 324).

It may be said here d pro])os to these remarks that
the subvertebral caudal muscles are, as a rule, but feebly
developed in the higher groups of birds, unless some
special habit of the form demands them, as we know
to be the case in the AVoodpcckers.

The following muscles in the trunk of the Raven
present themselves for our examination : — -

123. The complexus. 125. The flexor capitis inferior.

124. The rectus capitis anticus 126. The rectus capitis posticus

minor. major.

THE MUSCULATURE OF THE TRUNK.

263

127. The biventer cervicis.

128. The longus colli posticus.

129. The sacro-lumbalis.

130. The longissimus dorsi.

131. The obliquus colli.

132. The longus colli anterior.

1 33. The rectus capitis lateralis.

134. The trachelo-mastoideus.

135. The interspinales.

136. The interarticulares.

137. The obliquotransversales.

138. The intertransversales.

139. The triangularis sterni.

140. Tbe intercostales.

141. The scalenus medius.

142. The levatores costarum.

143. The appendico-costales.

144. The obliquus externus ab-

dominis.

145. The obliquus internus ab-

dominis.

146. The rectus abdominis.

147. The transversalis abdomi-

nis.

148. The diaphragm.

149. The levator coccygis.

150. The levator caudse.

151. The transversus perinei.

152. The depressor caudfe.

153. The depressor coccygis.

154. The lateralis caudal.

155. The lateralis coccygis.

156. The infracoccygis.

123. The complexus^ in the Raven, as in tlie vast

^ For the following synonymy and remarks I am indebted to
Gadow (loc. cit., p. 109) : — •

" 6. M. COMPLEXUS.

Complexus. Vicq d'Azyr, 1773, p. 581, No. 5.

Der Kopfdreher. Merrem.

Grand complexus. Cuvier ; Gervais et Alix, p. 15.

Der durchflochtere Muskel. Meckel, System, p. 297, No. 2.

Bauschmuskel des Kopfes (Splenius). Wiedemann, p. 75.

Bauschahnlicher Muskel. Tiedemann, p. 282.

Complexus. d' Alton, p. 7.

„ Owen, Cyclopcedia, p. 291.

„ Selenka, p. 98, No. 14.

„ Watson, p. 62.

" Dieser Muskel gehort mit dem M. biventer zur oberflachlichen,
selbststandig gewordenen Schicht des M. semispinalis in der oberen
Halsregion. Er entspiingt bei Colyvibus sehnig von der Endsehne
des M. semispinalis cervicis in Hobe des 5 Halswirbels, mehr fleischig
von den Intertransversarii posteriores und mit einem kurzen beson
deren Zacken von den Proc. ohliqui des 4 und 3 Wirbels. Sein

264 THE MYOLOGY OF THE RAVEN.

majority of the class, is a very distinct and well-
developed muscle.

It is clearly illustrated in my Fig. 70, and its mode
of orio;in and insertion in Fi^s. 4 and 69.

On either side of the neck, it arises 1;)y three tendinous
slips, one each coming off from the fourth, fifth, and sixth
cervical vertebrae, respectively. These several origins
occur upon the transverse processes of these vertebrae,
upon the supero-external aspects of the outer walls of
the lateral canals (Fig. 68). The slips pass between the
muscles of the region there found ; and, becoming
carneous, unite externally to form a broad sheet of
inuscular tissue, which, being superficial and overlying
the muscles of the occipital region for the most part,
passes round to meet in the median line the fellow of
the opposite side, in a tough fascia, forming a raphe
nearly two centimetres long. The two muscles thus
blended are now inserted into the occiput, some two
millimetres above the occipital ridge, as a thin tendinous

XJrsprung liegt demnach zwischen dem M. longus lateralis iind dem
M. semisjnnalis cervicis. Der Muskel wird ziemlich breit und stark,
wird nur vom M. cutaneus colli bedeckt, wahrend er selbst den M.
rechis cajntis 2)osticus nebst der Insertion des Jf. biventer uberdeckt.
Er inserirt sich an der queren Crista des oberen Eandes der Occi-
pitalia, in der Mittellinie mit dem der anderen Seite zusammen-
stcssend.

" Bei den mei.sten /Spheniscidoi entspringt er von den Processiis
ohliqui ]yosteriores des 5-3. Wirbels, bisweilen jedoch ist er auf zwei
Wirbel beschiankt. Bei den meisten Yogelu kommt er vom 3 und
4 Halswirbel."

Note. — Gadow gives five coloured figures in liis plates showing the
mvxscles in tl>e neck of various birds, and in these the complexus in
the neck of Cohjmhus septentrionalis appears to be exactly as I
describe it for the Raven ; while in the figure of the same bird on
another plate it appears to have a somewhat different origin (com-
pare Taf. 18a, figs. 1 and 2; 18b, fig. 1).

THE MUSCULATURE OF THE TRUNK. 265

sheet, the most superficial structure of the kind there
found.

The semi-ligamentous fascia, connecting these two
muscles in the median line, and extending forwards
to become inserted into the occiput, seems to represent
the only thing that birds can claim as the analogue
of a ligamentum nuchcB. In man, it will be remembered,
this important ligament is in the line of union between
the two trapezii muscles, and passes between the
neural spine of the seventh cervical vertebra and the
mid-point of the " su23erior curved line " of the supra-
occipital bone.

In Cormorants and the Anhinga a free bony " nuchal
style" is found j)rojecting from a mid-point upon
the occiput, in the convexity upon either side of
which a temporal muscle becomes inserted. I am
under the impression that I have said elsewhere in
my writings, and Coues has made the same statement
{Key, second edition, pp. 723, 724), that this bony style
of the Cormorants lies in the line of the analogue of
the ligamentum nuchcB in birds. But it is evident
that it cannot safely be compared in either case
with the ligament in question as it occurs in the
Mammalia. In short, in view of the fact that there is
no evident necessity whatever for the development of
such a support to the head in Aves, I must believe
that the wisest step to adopt in the premises is to
deny the occurrence of the ligamentum nucIicB, in
their class, altogether.

124, The rectus capitis anticus minor is a muscle in
the fore part of the neck in most birds, wdiicli has thus
been named and described by both. Gurlt and Owen,
And if I be correct in my diagnosis of it in Corvus, I
find it to arise in the Raven from the apices of the

266 THE MYOLOGY OF THE RAVEN.

hyapophyses of the second, tliird, and fourtli cervical
vertebrae, and winds round the Lateral aspect of the neck
in precisely the same manner as I descriljed for the
comp)lexus ; indeed, the opposed edges of these two
muscles are closely connected by fascia as they reach
each other, after which the present muscle passes up with
it to become inserted along on the same continuous line
just above the occipital ridge, and behind the bony
meatus auditorius of the skull.

The tendinous slips of origin of the rectus capitis
anticus minor pass between the Jlexor cajntis inferioris
and the longus colli anterior, being intimately con-
nected with the latter, and arising in common with the
former.^

^ We obtain the following in reference to this muscle from
Gadow {loc. cit. p. 120) :—

"16. M. RECTUS CAPITIS ANTICUS MINOR S. LATERALIS,

Le muscle droit (et moyen). Vicq d'Azyr, 1773, p. 582, No. 5.
Kopfbieger (pt.). Merrem.
Langer Kopfbeuger (pt.). Wiedemann, p. 76.
Flexor longus capitis. Tiedemann, § 210.

Vorderer, kleiner, gerader Kopfmuskel. Meckel, System, p. 299, No. 4.
Rectus capitis anticus minor. Gurlt, p. 17 ; Owen.
Trachelo-mastoideus. Owen, Apteryx, p. 285.
„ Selenka, p. 100, No. 20.

„ „ Watson, p. 64.

Occijnto-sous-cervical. Gervais et Alix, p. 15.
Occipito-sous-vertebral. Alix, p. 382.

'* Nicht so kraftig wie der ihn bedeckende Rectus major. Er setzt
sich in der Regel aus drei bis vier deutlichen Theilen zusammen,
die von der Ventralflache des 2 bis 6 Wirbels entspringen, und
sich alia zu einem Muskelbauche vereinigen, der mit einer kurzen,
rundlichen Sehne sich an dem eigenthiimlichen zitzenfiirmigen
Tuber der Seiten-Unterfljicho des Occipitale hasilare befestigt, dicht
hinter dem Meatus auditorius. Der betreffende Fortsatz ist nicht
mit dem Proc. Mastoideus zu verwechseln, er konnte als Proc.
paramastoideus bezeichnet werden."

THE MUSCULATURE OF THE TRUNK. 267

125. The flexor capitis inferior^ (Figs. 3, 4, and 67)
might almost be described as an " azygos muscle," so
intimately blended are the two in the median line.

It arises in common with the rectus capitis anticus
minor from the apices of the hypapophyses of the second,
third, and fourth cervical vertebrae, and in the pre-
sent instance either the fascia or a delicate tendinous
slip may even come off from the corresponding point
on the atlas. The thin tendons of origin of this muscle
soon become carneous as they pass forwards in some-
what of a bulky mass, uniting with the fellow of the
opposite side, to their insertion. This latter occupies
the entire triangular area offered by the basitemporal
of the base of the cranium, the bounding lines of the
muscle taking form therefrom, and in accordance there-
with.

This muscle in its action is a direct flexor of the
head upon the neck, whereas the rectus capitis anticus

1 As a rule this muscle has been considered to be one of the recti
muscles rather than a flexor of the head upon the neck which I
really conceive it to be. Its synonyms are tlius recorded by Gadow
{loc. cit., p. 120) :—

"15. M. RECTUS CAPITIS ANTICUS MAJOR S. MEDIALIS.

Le muscle (droit et) moyen de la tele. Vicq d'Azyr, 1773, p. 582,

No. 5.
Kopfbieger (pt.). Merrem.
Langer Kopfbeuger (pt.). Wiedemann, p. 76.

Mittlerer Kopfbeuger (Flexor medius capitis). Tiedemann, § 209.
Droit anterieur. Cuvier.

„ „ Gervais et Alix, p. 15.

Alix, p. 382.
Rectus capitis anticus major. Gurlt, p. 17 : Owen, Proc. Zool. Soc,

1842, p. 29 ; Watson, p. 64.
Recti capitis antici {major et minor). Selenka, p. 100, No. 20.
Vorderer, grosser, gerader Kopfmuskel. Meckel, System, p. 299,

No. 3."

2G8 THE MYOLOGY OF THE RAVEN.

minor ratlicr pulls it downwards, while tlic com-
plexus, no doubt, is an auxiliary to the more powerful
extensors found at the back of the neck.

The single carotid artery becomes nearly superficial
in the median line, near the posterior apex of the pre-
sent muscle, formed by the union of the two inferior
flexors of the head, just described, being covered only
by the nearly transparent common envelope of fascia.

It bifurcates soon after its emergence from the point
in cpiestion, and each lateral branch is harboured in
the sulcus formed l)y the present muscle and the rectus
capitis cmticus nu7ioi% until the vessel arrives at that
foramen which affords it, on either side, ingress to
the cranial cavity,

126. Tlie rectus capitis p)osticus rtiajor^ is, on either

^ The only muscle I can find in literature wherewith to compare
the present one is the rect. cap. posticus of Gadow, who presents us
with the subjoined synonymy for it, and in Bronn's Klassen des Thier-
Keiclis [loc. cit., p. 112) gives a description for it : —

" 8. M. RECTUS CAPITIS POSTICUS.

Kleiner Kopfheber. Merrem.

Le 2)etit viuscle droit posterieur ; petit complexu^. Cuviei'.
Rectus ccqjitis posticus major et minor. Tiodemann, § 205-^206.

,, „ „ „ „ „ d' Alton, p. 8.

Ohne Namen. Meckel, ISystem, p. 298.
Recti capitis jMstici. Selenka, p. 99, No. 16.
Grand et petit droit posterieur. Gervais et AHx, p. 14.
Rectus capitis p)osticus minor. Watson, p. 63.
Splenius cajntis. Watson, p. 63."

Note. — The muscle is by no means a small one in the Raven,
that is if I am correst in comparing my rect. cap. p)Ost. major with
the above one of Gadow and other authors. A comparison of the
synonyms of the recti muscles throughout the class Aves is
sufficient to convince anyone of the fact that they still stand in
need of careful diagnosis ; and further comparative reseaich in
representatives of divers groups of birds is a thing much to be
desired, in so far as their myology goes.

270 THE MYOLOGY OF THE RAVEN.

side, a powerfully developed muscle, arising, some-
what tendinous, in the median line, upon the anterior
aspect of the second cervical vertebra, all the way
from the summit of its neural spine to the mid-
point of the supero-anterior l)order of its neural canal.
The stronff bundle of carneous fibres formino- the
muscle diverge as they proceed to the occiput from this
linear origin, and are inserted upon a crescentic area
at the back of the skull. This insertion is within the
occipital line, being convex outwards, and situated well
towards the mastoidal aspect of the cranium (Fig. 4).

Evidently the united action of these muscles will tend
to extend the head upon the neck, and cither of them
acting independently will tend to pull the head towards
that side to which the contracting muscle belono^s.

127. Tlie hiventer cei'vicis ^ of Wiedemann is an ex-
ceedingly interesting muscle, or rather pair of muscles,
though they are well separated from each other. Upon
looking at the back of the neck of a Raven from

1 " M. BIVENTER OERVICIS.

Secundum par colli. Aldrovandi.
Digastricus. Steno.
Diinner Halsmuskel. Merrem.
Biventer cervicis. Wiedemann, p. 75.

,, ,, Tiedemann, § 203.

„ „ d' Alton (Zvveibauchiger Nackenmuskel), p. 8,

No. 2.

„ „ Gurlt, p. 17.

,, „ Selenka, p. 95, No. 5.

„ ,, Watson, p. 61.

„ „ Meckel, System, p. 295, No. 1.

Digastrique du cou. Cuvier.
Longus colli jMsticus (pt.). Owen.

Lo7ig posterieur du cou (^faisceau occipitcd) . Gervais et Alix, p. 1 4.
Faisceau interne ou digastrique du long interejnneux cervical. Alix,
p. 379." (Gadow in Bronn's Klassen, loc. cit., p. 107.)

THE MUSCULATURE OF THE TRUNK. 271

wliicli the integuments have been removed, we notice
at about its middle a deep, longitudinal cleft, which
becomes shallower as it approaches its ends, and
finally flush with the general muscular surface of the
neck. The lateral borders of this cleft, at their
middles, show, beneath the enveloping and thin fascia,
each a longitudinal tendon. These are the mid-tendons
of the muscles we have now under consideration.

A hiventer cervicis arises by a slight tendon from the
side of the neural spine of the first dorsal vertebra,
and from the neighbouring tendon of the longus colli
posticus muscle. This flattened tendon passes directly
up the back of the neck, closely applied against the last-
named muscle.

At the end of rather more than two centimetres it
terminates in a flat spindle-shaped muscle, the tendon
beino- ao;ain resumed from the anterior end of the
same. This latter division of the tendon is that por-
tion which bounds the cleft above described, passing
which, once more a flat muscle is developed, which over-
lies the longus colli posticus anteriorly, and terminates
by being inserted into the occiput, to the outer side
of the occipital prominence, and between the inner
extremities of the insertions of the complexus and
rectus capitis posticus major (Fig. 4).

In speaking of this muscle in the Apteryx, Owen
says : " A slender fasciculus is detached from the mesial
and dorsal margin of the longus colli posticus, near
the base of the neck, which soon terminates in a
long round tendon ; this tendon is traced down by
short aponeurotic fibres to the s^^ine of the fifth,
fourth, third, and second cervical vertebrae inclusive,
immediately beyond which it again becomes fleshy,
and expands to be inserted into the occipital ridge ;

:^72 THE MYOLOGY OF THE RAYEN.

tliis portion is the digastrique or hiventer capitis of
Cuvier" {Anat. of Verts., vol. ii. p. 88).

These muscles are also present in the Eaptorial hirds.
128. The longus colli posticus is, on either side, a com-
plicated muscle extending the entire length of the back
of the neck, from its base at the anterior portion of the
dorsum, to its final insertion upon the vertehra dentata.
It arises by a thin sheet of tendon from the mar-
ginal edges of the summits of the neural spines of
the first two leading dorsal vertebrae. This sheet of
tendon appears to be in continuation with the tendon
of orio"in of the longissimus dorsi, and like it is irregu-
larly split up into narrow little ril)bonlets of varying
wddth, and some five or six in numl)er.

This tendinous origin of the longus colli posticus soon
becomes muscular as it advances up the neck, and as a
lono-, somewhat narrow, flattened muscle it extends the
entire length of this part, to become finally inserted
into the transverse process of the axis vertebra.

From the under side of this muscle, at its dorsal
extremity, it throws down certain fleshy fasciculi. The
first or most posterior one of these blends with the
muscular portion of the longissimus dorsi. Then
follow six well-defined slips, each flattened from before
backwards, and each becoming narrower as we proceed
towards the head. The most anterior fasciculus of this
series is the longest, and they progressively become
shorter as we proceed towards the thorax.

Eeo'ardinff the neck from a lateral view, and liftino^
up the longus colli posticus, we observe that these
descending fasciculi pass obliquely from the under
side of the muscle forwards to their insertions. The
first of these latter is the postzygapophysis of that
cervical vertebra which supports the shorter pair of

THE MUSCULATURE OF THE TRUNK. 273

free cervical ril;)s. The remaining five fasciculi makes
similar insertions, but become more and more intimately
blended with the ohliquus colli muscles of the same side,
and which make similar insertions.

Now, where these fasciculi terminate in the middle
of the neck, wc find another series commencino-. Callinsi:
the one at the middle of the neck the first of this
new set, we find it to be a long slender slip which
arises, tendinous, from the neural spine of the seventh
cervical vertebra, and extending obliquely forwards
(just the reverse of the last series) it merges with
the fibres of the under side of the longus colli i?osticus
proper. The next in order of this series of fasciculi is
shorter and thicker, and thus they proceed until the
last or sixth one, coming from the neural spine of the
axis vertebra, blends very intimately with the inner
margin of the longus colli posticus at its proper inser-
tion. This last fasciculus is the shortest and thickest
of this series.

Professor Garrod gave an excellent figure {P.Z.S.,
Plate xxvi.) of the very interesting arrangement of the
longus colli 2'>osticus in the Plotus anhinga, and after
describing its peculiarities as they are found in that
l)ird, he says, in conclusion, that " It is nearly always
the case in avian anatomy that the inner fibres of
the cervical portion of the longus colli 'posticus muscle
become difi'erentiated to form the cligastrique du cou
of Cuvier, better known to us as the hivenfer cervicis,
a muscle one peculiarly interesting modification of
which, in the genus Ceryle among the Alcedinidse, has
been described and figured by Dr. Cunningham in the
Society's Proceedings (1870, p. 280). This, by the way
I may mention, I have had the opportunity of fully veri-
fying, Meckel, in his General T'reatise on Comparative

T

lvve,nte/i^ c^rvtci^.

Fig. 70. — Right lateral view, life-size, of the head and neck of a Eaven, dissected
and drawn by the author. Designed to show the siiperHcial muscles of
the region. Those at the back of the neck are lifted from their posi-
tions ; while the bivcnter cervicis and compUxus are widely separated in
order to show the muscles lying beneath them.

THE MUSCULATURE OF THE TRUNK. 275

Anatoyny, tells us that lie found it at its minimum of de-
velopment in the Gallinae, the Goose, and the Cormorant.
In a specimen of Sulafusca, as well as in Phalacrocorax
carho, it is present, but extremely small, I find. It is
entirely absent in Plotus anhinga, the longiis colli
posterior {cervicalis ascendens, Meckel) entirely ceasing
at the lower margin of the axis vertebra, in the tendon
above described."

The longus colli jwsticKS in the Ajyteryx, as described
by Professor Owen, differs in the number of fasciculi
of its accessory series, and other minor details, while
in the main it practically agrees with the muscle as
I have here described it for the Raven. I regret to
say that Meckel's work is not at present at hand,
and consequently I am debarred from making com-
parisons that would greatly enhance the value of my
work.^

^ I here present Owen's description of this mviscle in the AjJteryx,
as it may be of interest to compare it. He says: — " The longus colli
2)ostiGus is most internal or medial of the superficial muscles of the
dorsal aspect of the thoracic and cervical regions. At its posterior
part it seems to be a continuation of the longissimus dorsi ; its
medial and anterior part offers a strong analogy with the hiventer
cervicis ; it is the homologue of the first, or medio-dorsal series of
the oblique fibres of the muscular system in fishes. It commences
by long and slender, but strong, subcompressed tendons from the
spines of the sixth, fifth, and fourth dorsal vertebrae : these tendons
gradually expand as they proceed forward and downward, and send
off from their under surface muscular fibres which continue in the
same course, and begin to be grouped into distinct fasciculi at the
base of the neck ; the first of these bundles joins a fasciculus of the
longissimus dorsi, which is inserted into the anapophysis of the
thirteenth cervical vertebra ; the succeeding fasciculi derive their
origins from a broad and strong aponeurotic sheet attached to
the spines of the fourth, third, and second dorsal vertebrae : the
second to the eighth fasciculi inclusive are compressed, broad,
and fieshy, and are inserted in the strong round tendons de-

T 2

276 THE MYOLOGY OF THE EAVEN.

129. The sacro-lumhcdis muscle in the Raven is but
moderately developed, and quite intimately l)lended
with the longissinius dorsi throughout its entire
extent.

It is only in the highly organized vertel^rates (Homo)
that we find a great complexity of the musculature of
the dorsal region. Mivart, alluding to this matter, says
that " the erector S'pinfB presents in man a degree of
differentiation not generally found in animals below
his class. Thus in the Iguana and Chameleon it is but
divisible into the longitudinal parts answering respect-
scribed in the preceding miiscle [ohliqutis ohliq'td], and attached to
the zygapophysis of the twelfth to the sixth cervical vertebrte
inclusive : the ninth fasciculus, which forms the main anterior
continuation of the longtis colli 2^osticus, is longer than the rest,
and receives, as it advances, accessory fibres from the spinous
processes of the seventh to the third cervical vertebrae inclusive, and
is inserted, partly fleshy, partly by a strong tendon, into the side of
the broad spine of the vertebra dentata " {Anat. of Verts., vol. ii.
pp. 87-88).

It is three years since this footnote was written, and at this date
I can add to it the synonymy of the longvs colli posticus as compiled
for us by Gadow, who designates the muscle as his " System des M.
spinalis." It is as follows : —

" 7. System des M. spinalis.
S^nnalis dorsi. Gurlt, p. 18.

,, ,, Selenka, p. 96, No. 6.

Long posterieur du cou (pt.). Gervais et Alix, ]). 14.
Longissimus dorsi (pt.). Watson, p. 56.

"Theil III.
Strecker des Tragers. Wiedemann, p. 76.
M. extensor atlantis. Tiedemann, § 212.
Halsdornmuskel. Meckel, System, p. 294, No. 5.
Longus colli p>osticus (pt.). Selenka, p. 95, No. 4.
Longus colli posterior. Garrod, Proc. Zool. Soc, 1876, p. 338.
Extensor magnus colli (pt.) ; sjjlenius colli. Watson, p. 57." (Bronn's
Klassen des Thier-Reichs, vi. Bd., p. 110).

THE MUSCULATURE OF THE TRUNK. 277

ively to tlie longissimus dorsi and sacro-lumhalis, and
continuing, with the intervention of certain neck muscles,
from the cranium to the end of the dorsum of the taiL
But a great simplicity still may exist, as in Tailed Batra-
chians (e.g. Menopoma and Menohranclius) , where, with-
out the intervention of any such neck muscles, a simjjle,
or more or less tendinously intersected muscular mass
extends from the skull directly to the end of the dorsum
of the tail. This dorsal muscle may be reduced to a
mere rudiment, as in Emys, where it runs between the
transverse and neural processes and the carapace "
{Elem. Anat, pp. 322, 323).

Among the group of muscles we are now examining,
the deep layer of muscles of the back in the Eaven, the
sacro-lumhalis forms the outer j^ortion of that close-
fitting, tendo-muscular sheet seen extending between
the anterior margin of the ilium to the root of the neck,
and is to be only with difficulty differentiated from the
longissimus dorsi, which is to be found between it and
the dorsal neural spines.

It arises,^ quite tendinous, from the anterior margin

^ The system of musculature of the dorsal region in birds, Professor
Gadow defines as the " I. Dorso-spinale Muskeln," and proposes
to divide this " System des M. sacro-spinalis " into four parts, viz.
— 1. M. iLiocosTALis ; 2. M. longissimus dorsi; 3. M. cervicalis
ASOENDENS ; and 4. M. transverso-oeliquus. Of these, the first
represents the muscle we now have under consideration, or the
SACRO-LUMBALis, and for it Gadow sees the following synonymy,
viz. : —

"1. M. ILIOCOSTALIS.

Costo-cervical (pt.). Yicq d'Azyr, 1772, p. 580, No. 2.

Aeusserer Riickgrats-Strecker. Tiedemann, § 217.

Der Riickenmuskel. Merrem.

Opisthotenar (Aeusserer Bauch). Meckel, System, p. 291, ISTo. 1.

Sacro-lumhalis. Owen; Gurlt, p. 18

278 THE MYOLOGY OF THE KAVEN.

of the ilium ; from tlie angles of the last two vertebral
ribs ; and by tendinous slips from the outer ends of
the transverse processes of the last three dorsal
vertebrae. These last tendinous fasciculi spread out
upon, and merge with, the under side of the muscle, and
probably represent, as Owen says, the musculi accessorii
ad sacro-lumbalem. To be seen, the free and outer
margin of the muscle under discussion must be elevated.

O

The sacro-hmibalis is inserted by a few fleshy fibres
into the angle of the first dorsal rib, and in muscular
subjects sometimes by a few additional fibres to corre-
sponding 2^oiiits upon the free cervical ribs ; while its
main insertion is by a strong semi-tendinous insertion
into the outer extremity of the diapophysis of the twelfth
cervical vertebra.

As we might have expected. Sir Eichard Owen
found the insertion of this muscle consideraljly more
extensive than this in the Apteryx, and this eminent
anatomist seemed to see in the final insertions of the
sacro-lumbalis in that bird, the representatives of the
cervicalis descendens and ascendens as they have been
described for man.

130. The longissimus dorsi ^ has already been alluded

Sacro-lumhalis. Selenka, p. 93, No. 1.
,, ,, Gervais et Alix, p. 13.

,, „ Watson, p. 55."

Note.- — For Gadow's descriptions of his divisions of the spinal
system of muscles,' see Bronn's Klassen des Thier-Eeichs, vi. Bd., pp.
105-107.

^ See footnotes under the sacro-lumhalis muscle of the present
work (No. 129), and the following synonymy will be made clear: —

" 2. M. LONGISSIAIUS DORSI.

Costo-cervical (pt.). Vicq d'Azyr.

Innerer Ruckgrats-Strecker (sacro hwibalis). Tiedemann, § 217.

THE MUSCULATURE OF THE TRUNK. 279

to, and its relative position defined in describing
the last muscle. In Figs 66 and 67, I find that I
can indicate only the principal origins and insertions
of these less important muscles of the back, as an
injury would be done the drawings, and its general
clearness interfered with, if all the smaller origins and
insertions were given, together with the necessary
lines to indicate them.

The present muscle has quite a complicated origin,
though its insertion is rather simple. It arises from
the inner moiety of the anterior margin of the ilium,
and from the various surfaces afforded it by the walls
of the corresponding " ilio-neural canal," or that channel
existing between the ilium and the crista of the sacrum
in front. In some specimens I have seen the posterior
tendinous ends of the longissimus dorsi extend beyond
the hinder opening of this channel, and in some cases
almost reach the superior caudal muscles. Of course,
in Eeptiles, the muscle is continuous the entire length
of the back and tail, to the very tip of the latter.

In our subject the longissimus dorsi also arises by
a series of short and distinct tendons alternately from
the anterior and posterior extremities of the summits
of the neural spines of all the dorsal vertebra3.
It will be remembered that the free margins of these
neural spines above are l^ifurcated behind and pointed
in front. Now the tendons of this series attached to
the points in front extend forwards and merge into
the muscle, and are tendons of insertion, while those

Opisthotenar (Tnnerer Bauch). Meckel, System, p. 291, No. 1.
Longissimus dorsi. Giirlt, p. 18 ; Owen.

,, „ Selenka, p. 94, No. 2.

„ „ Watson, p. 56.

Long du dos. Gervais et Alix, p. 13."

280

THE MYOLOGY OF THE RAVEN.

^nte^T-spi^iaZ^

^<57- 72.

Fig. 71. — The upper figure is a right lateral view of the first six cervical vertebrae,
including tlie axis and atlas (size, x l^). a points to that portion of
tile infrHransvcr.'ialis muscle wliicli passes between the parapophyscs of
the fifth and sixth cervicals. (See description in text.)

Fig. 72. — Dorsal view of the sixth, seventh, and eighth cervical vertebraj ( x 2),
showing representations of three other sets of deep muscles of this
region — the intertransversalcs, the inter articular es, and obliqno-trans-
vcrsalcs. Drawn by the author, from his dissections of the Kaven.

coming off from the tip of the bifurcation behind take
ju,st the opposite course, and are additional tendons of
origin. So when we pull the muscle away from the

THE MUSCULATURE OF THE TRUNK. 281

sides of the dorsal neural crests, a system of X^ are
seen to be formed by these little tendons crossing each
other.

The longissimus dorsi in the Eaven also arises, some-
what fleshy, from the diapophyses of the dorsal
vertebrae, and from the fascia between it and the
sacro-lunibalis , and also by a tendinous sheet continu-
ous w^ith the origin of the longus colli 'posticus from
the free surfaces of the crests of the neural spines of
the last three dorsal vertebras.

It will be seen from all this, that the present muscle,
on either side, fills in the " ilio-neural canal " of the
pelvis ; the open angle formed by the neural spines and
diapophyses of the dorsal vertebrae ; and is bounded
externally principally by the sacro-lumhalis muscle.

Now the longissimus dorsi is further inserted by four
fasciculi, the anterior one being strong and tendinous,
into the free hinder margins of the oblique j)rocesses
of the eleventh, twelfth, thirteenth, and fourteenth
vertebrae, respectively.

These insertions are so disposed that the ohliquus
colli, or the series of muscles so called, seem to con-
stitute the harmonious continuation of them up the
side of the neck ; while above, the longissimus dorsi is
apparently so continued up the back of the neck by
the longus colli jwsticus.

131. The ohliquus colli ^ consists of a series of

^ Turning to the sacro-lumhalis (No. 129) of the present work,
and consulting the footnotes there given, the following synonymy
is further illustrative : —

" 4. M. TKANSVERSO-OBLIQUUS.

Extensor parvus colli. Wiedemann, p. 77 ; Tiedemann, § 214 ;

Watson, p. 58.
Quermuskel des Nackens. Meckel, System, p. 294, ISTo. 5.

282 THE MYOLOGY OF THE RAVEN.

seven oblique fasciculi of muscles seen at the side of
tlie neck. The first or posterior one of these is the one
which links the continuation of the system commenced
by the longus colli 2'>ostic'US, the longissimus dorsi, and
the sacro-lumhalis, and which is continued by the re-
maining fasciculi of the present muscle.

This first or posterior representation of this com-
plicated series, as I say, arises from the diapophysis
of the eleventh cervical vertebra, winds obliquely over
the tenth vertebra to become inserted into the hinder
margin of the postzygapophysis of the ninth vertebra,

Ohliqu^is colli. Owen.

Ohliquo-transver sales. Selenka, p. 97, No. 11."

In order to make the spinal system of muscles in birds complete
as worked out by Gadow, we must yet quote his third part, which
is as follows : — ■

" 3. M. CERVICALIS ASCENDENS.

Long exte^iseur du cou (pt.). Yicq d'Azyr.

Grosser Halsstrecker (pt.). Tiedemann, § 213 ; Wiedemann, p. 76.

Longus colli 2)ostic^is (pt.). Owen; Selenka, p. 95, No. 4.

Aufsteigender Nackenmuskel (pt.). Meckel, Sysfem, p. 293, No. 1.

Cervicalis descendens. Gurlt, p. 18.

Long posterieur du cou. Gervais et Alix, p. 1 4.

Extensor magnus colli (pt.). Watson, p. 57."

Note. — In consulting this synonymy, it must be compared with
the account of the longus colli jwsticus muscle of the present work,
of which Owen considered the above muscle to be but a part : —

" 3. M. CERVICALIS ASCENDENS.

" Der laterale Theil {Cervicalis ascendens). Seine oberfliichlichen
Fasern sind directs Fortsetzungen des M. iliocostcdis, oder der seit-
lichen Theile des M. longissimus dorsi. Er entspringt in seinem
hintersten (caudalwarts) Theile von den Rippen, hauptsachlich aber
von der Oberflache der Querfortsatze, weiter kopf wiirts aber von den
Enden oder Spitzen derselben. In der ganzen Liinge des Halses ist
der Cervicalis ascendens mit der unteren seitlichen Halsmuskulatur
verwachsen, die an der unteren Kante der Querfortsatze und an den
Halsrippen entspringt." (Gadow, loc. cit., p. 107.)

THE MUSCULATURE OF THE TRUNK. 283

being joined in its passage by the third descending
fiisciculus from the longus colli posticus (the fasciculi
ohliqui of the longus colli posticus, or rather the third
one of the series as so described by Owen ?).

Now, the next fascicukis of the muscle we have
under consideration arises from the transverse process
of the tenth cervical vertebra, winds aljout, parallel
with the last, to skip the ninth, and be inserted into
the hinder margin of the postzygapophysis of the
eighth vertebra ; it also being joined by the next
fasciculus in order descending from the longus colli
posticus.

The next three fasciculi arise in a similar way, but
in addition to being attached to the postzygapo-
physes they continue forwards to make an insertion
also upon the neural spines of the alternate vertebrae
in turn, being inserted in fact with the anterior set of
descending fasciculi from the longus colli posticus.

The insertion of the last two slij^s of the ohliquus
colli again differ. The most posterior of these two
arises from the transverse process of the sixth vertebra,
and passing less obliquely up the neck, becomes in-
serted into the extremity of the cliapophysis of the
fourth vertebra, by a strong tendon ; the outermost
carneous fibres of the fasciculus, still continuing towards
the head, make another tendinous attachment to the
outer extremity of the transverse process of the third
vertebra.

Now the last or most anterior fasciculus of this series
arises from the transverse process of the fifth vertebra,
and passing under and parallel with the last, is in-
serted with its leading tendon into the extremity of
the diapophysis of the third vertebra. This completes
one of the most complicated systems of muscles that

284 THE MYOLOGY OF THE RAVEN.

I have any knowledge of, and is scarcely excelled by
the so-called, and famons " fifth layer " that adorns the
dorsal region of Homo.

Owen fonud the ohliquus colli fully developed in
the Ajyferi/x, and says of it that " this series of muscles
seems to represent the transversalis colli, wdiicli is
the anterior continuation of the longissimus dorsi
in Mammalia, but it differs in being inserted into the
oblique, instead of the transverse processes. In the
direction of their filjres these fasciculi resemljle the
semis'pinalis colli, l:)ut they are inserted into the
oblique processes instead of the spines of the vertebrae "
(Anat. of Verts., vol. ii. p. 86). It will be noticed that
we found in the middle of the series in our j^resent
sul)ject that they do pass to the neural spines.^

1 In his dissections of the Ajjteryx, Professor Owen also made out
in the dorsal region of the back the spinalis dorsi, the multijidus
S2n7ice, and the ohliquo-spinales. These muscles in the Raven cannot
be satisfactorily differentiated — if indeed they are present at all — or
even their barest rudiments. Upon a number of specimens I have
taken great pains to endeavour to isolate them, but was after all
forced to the conclusion that they do not exist in so high a type as
Corvus. I quote in full here from the second volume of the Ana'omy
of Vertebrates what Professor Owen says of them as they occur in
the Apteryx, so we may have the data for comparison. According
to this authority, the spdnalis dorsi is brought into view by the
removal of the dorsal portion of the longus colli ^^osticus and the
longissimus dorsi.

" It arises by two long, narrow, flattened tendons from the
spines of the eighth and seventh dorsal vertebrse : these pass
obliquely downward and forward, expanding as they proceed,
and terminate in two fasciculi of muscular fibres : the posterior
bundle passes forward beneath the anterior one, and inclining
inward and upward, divides into two portions, inserted by long-
tendons into the spines of the second and first dorsal vertebraj ; it
then sends a few fibres forward to join the outer and anterior
fasciculus, which is partly inserted by a slender tendon into the

THE MUSCULATURE OF THE TRUNK. 285

132. The longus colli anterioi- is that complicated
muse alar mass which is found runniiio- the entire

o

length of the anterior aspect of the neck, superficially.
To be properly studied it must first be carefully un-

spine of the last cervical vertebra : the rest of the fibres of the
second fasciculus join the portion of the longissimus dorsi which is
implanted into the posterior oblique process of the last cervical
vertebra. The three inserted tendons of the spinalis dorsi are also
the medium of attachment of fibres continued from the muUifidus
s2)incG, beneath them.

" The series of muscles called multifidus spince arises by fleshy
fibres from the diapophyses of the five last dorsal vertebrte, which
pass upward, forward, and inward, to be inserted by four flat tendons
into the spines of the seventh to the third dorsal vertebrae inclusive,
and by the tendons of the spinalis dorsi into the two anterior dorsal
spines.

" Ohliquo-spinales. The removal of the multifidus spince brings
into view a series of long, narrow, flat tendons, coming off from the
spines of all the dorsal vertebrae, and slightly expanding as they
proceed forwards and obliquely downwards and outwards ; they
become fleshy half-way from their origin, and are inserted into the
posterior oblique and ti-ansverse processes of the six anterior dorsal
vertebra;, and into the posterior oblique processes of the three last
cervical vei-tebr^e " (pp. 88, 89).

Although these muscles are of no particular importance as com-
pared with far more constant elements of the muscular system, and
still less so with such muscles as can be effectively utilized in classi-
fication, still their study and investigation affords not a little
material of interest when we come to search for them as facts
simply to illustrate our comparative researches.

To this end they may be advantageously compared with the dorsal
muscles of jEchid^ia and Ornithorhynchus and the lower forms of
birds.

For still further information vipon this point see Gadow's (" 9. M.
MULTIFIDUS -fSEMisPiNALis") account in Bronn's Klassen des Thier-
Reichs, vi. Band, pp. 113, 114 ; also Selenka in the same work [M.
onultifidus spinm), p. 96, No. 7 ; likewise the works of Gurit (p. 18),
and Meckel's System, p. 292, No. 2, The subject demands more
extended research and comparison.

286 THE MYOLOGY OF THE RAVEN.

rcavelled from one end to the other, and its various
origins and insertions neatly cleaned down to the very
bone.

We then find the muscle composed of three distinct
portions, as in some of the Mammalia, viz, a vertical
2)0)'tioii, and a su])crior and an inferior oblique
portion. Besides these, there are certain accessory
fasciculi to be examined.

The vertical portion of the longus colli is the major
division of the muscle. It arises, largely fleshy, from
the hypapophyses of the tenth to the fifteenth vertebra)
inclusive, and extends the entire length of the neck to
be inserted by a strong tendon into the inferior tubercle
of the atlas, alono-side of the similar tendon comino;
from the fellow of the opposite side, with which in this
locality it is intimately l)lended, — while attached to
the same tubercle to its outer side we find the tendon
of the superior oblique j)ortion. Now from the under
side of this division of the longus colli we find a
system of tendons given oft', wdiicli become longer and
longer as they reach up the column, and in turn be-
come attached to the apices of each and all of the
parapophyses of the cervical vertebrse to include the
tenth, and exclude the first three. It will be seen that
the hypapophyses of the dorsal vertebrae trifurcate, and
the part of the longus colli which arises from the
lateral processes thus formed, is more or less distinct,
especially posteriorly, but scarcely enough so as to
warrant its being described as a separate portion.

Mesially, at tlie anterior division of the neck, the
longus colli anterior, in our subject, also throws off
tendons as it passes the fourth, third, and second
cervical vertebraB, which respectively become attached
to the hypapophyses of these verteljrge. Beneath these

THE MUSCULATURE OF THE TRUNK. 287

tendons the carneous portion of the muscle is more or
less attached to the centra of the vertebrae in question,
which several attachments correspond to the accessory
fasciculi down the remainder of the neck to at last
distinctly include the eighth vertebra. Each of these
fasciculi arise from the fore part of the pleurapophysis
of a vertebra, and pass oblic[uely upwards to become
attached to the under side of that tendon of the
longus colli which is inserted into the apex of the
parapophysis of the vertebra next beyond.

This system of accessory fasciculi of the longus
colli anterior can only be satisfactorily examined after
the muscle has been forcibly pulled away from the verte-
bral column, catching ahold of it near its middle.

The superior ohlique portion of the muscle is quite
distinct, and arises from the diapophyses of the fifth,
fourth, and third vertebrae, and passing upwards becomes
tendinous, and is inserted, as already stated, into the
inferior tubercle of the atlas, to the outer side of the
tendon of the vertical portion.

The iyiferior oblique portion is also distinct, but must
be regarded more in the light of a specially differen-
tiated fasciculus from the vertical portion.

It arises from the transverse processes of the sixth,
fifth, and fourth vertebrae (mesiad to the preceding
portion on the fifth and fourth), passes upwards, and
is inserted by a delicate tendon into the apex of the
parapophysis of the third vertej^ra.

In the Proceedings of the Zoological Society of
London (1876, pi. xxvi.), Garrod gives an excellent
figure showing the interesting peculiarities of the
longus colli in Plotus, and Sir Eichard Owen says of
it in the Apteryx that " this large and long muscle,
which appears simple when first exposed, is found to

288 THE MYOLOGY OF THE EAVEN.

consist, when unravelled by further dissection, of a
series of closely succeeding, long, narrow fasciculi, arising
from the liypapophyses of the sixth dorsal to the first
dorsal, and from the ten posterior cervical vertebrae ;
and sending narrow tendons, which increase in length
as they are given off more anteriorly, obliquely for-
ward and outward, to l)e inserted into the pleura-
pophyses of all the cervical vertebra3 save the first
two : the highest or foremost tendon is attached to
the tubercle at the under part of the ring of the atlas ;
1)ut this tendon is also the medium of insertion of
five small fasciculi of muscular fil)res arisino- from the
diapophyses of the sixth, fifth, fourth; third, and second
cervical vertebrae" [Anat. of Verts., vol. ii. p. 90).

It will be seen here that this eminent anatomist
made no distinction of the superior oblique and inferior
ohlique portions ; and that the accessory fasciculi
apparently all go to the same tendon for their insertion,
whereas in the Eaven they pass to the several tendons
in turn, as they are thrown off from the main muscle
to become attached to the apices of the spine-like
parapophyses. It will be seen, too, that Professor
Owen says in the Apteryx that the tendons of this
muscle are attached to the pleurapophyses instead
of the parapophyses, as they arc in the Raven, and
as Garrod fio-ures them in Plotus}

o

^ Gadow's account of the longvs colli aiitic^is is very full and clear,
and of it he gives the following synonyms : —

" 14. M. LONGUS COLLI ANTICUS.

Long JiecMsseur du cou. Vicq d'Azyr, 1773, p. 582, No. 4.
Langer Halsbeuger. Wiedemann, p. 77.

„ „ Tiedemann, p. 291, No. 9.

Langer Halsmnskel. Meckel, p. 295, No. 1.
Longus colli. Gurlt, p. 19.

THE MUSCULATURE OF THE TRUNK. 289

133. The rectus capitis lateralis is a well-deve-
loped muscle in the present subject. It arises some-
what tendinous from the diapophyses of the fifth, fourth,
and third cervical vertel^rse ; and, passing obliquely
upwards in front of the spinal column, the fibres con-
verge to form a strong, sub-compressed tendon, which
is inserted into the inner tubercle on the basal ridge
of the basitemporal.

134. The trachelo-mastoideus^ is a powerfully deve-

Longus colli. Owen, Apteryx, p. 310.
Selenka, p. 100, No. 19.
,, ,, Watson, p. 60.
Long anterieur du cou.- Gervais et Alix, p. 15.
Longus colli anterior. Garrod, P.Z.S., 1876, p. 337.
Lmigus colli externus. Watson, p. 61." (The reader is referred to
Bronn's Klassen des Thier-lx'eichs, vi. Band, p. 118.)

^ No little confusion seems to exist in recognizing and dis-
tinguishing these remaining two cervical muscles, i.e. the rectus
capitis lateralis and the present one, the trachelo-mastoideus. For
my own part, I can show them no clearer, at present, than I have
attempted to do in Fig. 70 of this work. Even Gadow seems to
have seen the two muscles in his one that he has called the
" longus lateralis cervicis et capitis,'^ and of which he gives the
following account {loc. cit., pp. 116, 117) : — ■

" 13. M. LONGUS LATERALIS CERVICIS ET CAPITIS.

Les droits lateraux de la tete. Vicq d'Azyr, 1773, p. 582, No. 6,

Cuvier.
Grand transversaire. Cuvier (1).
Seitenbeuger des Kopfes. Wiedemann, p. 75.
Flexor capitis lateralis. Tiedemann, § 208.
Halsdorn-Muskel ; Quermuskel des Nackens. Meckel, System,

p. 294, No. 5.
Trachelo-mastoideus. Meckel (1).

,, „ Owen, Aj)teryx, p. 285 (X), Selenka, p. 99,

No. 15.
Rectus cap)itis lateralis. Owen, P.Z.S., 1842, p. 22.
Selenka, p. 99, No. 18.
„ „ „ Watson, p. 63 (pt.).

U

290 THE MYOLOGY OF THE HAVEN.

loped muscle, flattened from side to side, and situated
external to, and completely overlapping, the rectus
capitis lateralis.

It arises, semitendinous, from the diapophyses of
the fifth, fourth, third, and second cervical vertebrae,
and the fibres rapidly converging as they pass obliquely
towards the skull, unite to form a strong and sub-
compressed tendon, which is inserted into the outer
tubercle of the basal ridge of the basitemporal at the
base of the cranium.

This muscle apparently exactly agrees with the tra-
chelo-mastoideus in the Apteryx as described by Owen ;
except in this latter bird it seems to be inserted a

Basi-transversaire. Gervais et Alix, p. 15.
Longus colli externus. Watson, p. 61.

" Das specielle Verhalten der diesen Muskel zusammensetzenden
Blindel ist bei Anser wie folgt. Ein Biindel entspringt von der
unteren Halfte der Proc. transv. des 8 Wirbels, verbindet sich dann
mit der tieferen Partie des vom 7 Wirbel kommenden Biindels,
und inserirt sich am freien Ende der Halsrippen des 7 Wirbels.

" Die vom 7 Wirbel entspringenden Theile gehen zu den Hals-
rippen des 6 und 5 Wirbels,

die des 6 zu den Halsrippen des 5, 4 und 3 Wirbels,

5 4 .S "^

11 11 '-' ti 11 11 11 ^1 '-' 11 -J 11

4 3 2 1

11 11 ^ 11 11 11 11 'J, -J ,j J. .,

„ „ 3 und 2 „ „ „ „ 1 „

und zum hinteren Eande des Os occijntale basikxre, seitlich neben
dem Condylus.

" Es treten unter diesen Muskeln jedocli ausserst verschiedene
Spaltungen der Urspriinge und Verwachsungen der Insertionen
avif, sodass das oben gegebene Verhalten nur als ein schematisches
aufzufassen ist. Die Insertionen sind gewohnlich derart am
freien Ende der Halsrippen, dass haupsachlich die oberen Kanten
und hinteren Rander zur Befestigung dienen, wahrend die Seiten-
rander ziemlich frei bleiben, nur am 2 Wirbel und am Altas
riickeu die Insertionen entsprechend dem Fehlen des Halsrippen auf
die untere Seite der entsprechenden Fortsiitze."

THE MUSCULATURE OF THE TRUNK. 291

little more externally, as the authority quoted states
that its insertion is to be found on the paroccipital.

135. The inter sjnnales ^ constitute a series of muscles
that connect the neural spines of the vertebrae.

^ In Figs. 71 and 72 I have attempted to clearly portray the
system of muscles that may be considered to be strictly intervertebral
in the cervical region of the Raven, and they probably thus exist in
the vast majority of the class. Of the four sets, viz. the Inter-
sjnnales, the Interarticulares, the Ohliquo-transver sales, and the Inter-
transversales, Gadow gives good brief accounts, with their several
synonymies as he makes them out. All these latter I quote below,
with the exception of the Interarticulares, for which I fail to find
a description by the author in question, unless perchance he includes
them in his muscle No. 9, or the M. niultijidus + semisjnnalis, an
account of which I have quoted from him on a former page of
the present work (see Bronn's Klassen des Tliier-Reiclis, vi. Band,
pp. 113-115):—

"11. M. INTERSPINALES.

Zwischendorn-Muskeln. Meckel, System, p. 294, No. 6.

„ „ Tiedemann, p. 292, No. 12.

Obliquo-spinales. Selenka, p. 97, No. 8.
Inter sjnnales. Owen; Selenka, p. 97, No. 9.
Interarticulares. Selenka, p. 97, No. 9.
Court interejnneux. Alix, p. 374."

These I take to represent the Interspinales (No. 135), and the next
the Ohliquo-transver sales (No. 137), while the last are my Intertrans-
versales (No. 138) : —

" 10. Mm. kotatores s. obliquo-transversales.

Kleine Zwischenquermuskeln ('?). Meckel, System, p. 294, No. 7.

Midtifidus sjnnce. Owen.

Obliquo-transversales. Selenka, p. 97, No. 11.

Muscles articulo- ou epineux-transversaires. Alix, p. 378.

" 12. System dee Mm. intertransversarii.

Zwischen-Quermuskeln. Wiedemann, p. 78.

Vordere und hintere Zwischen-Quermuskeln (^Intertransversarii).

Tiedemann, § 221.
Intertransversaires. Cuvier, vol. i., p. 190.
„ Gervais et Alix, p. l4.

u 2

292 THE MYOLOGY OF THE RAVEN.

They do not occur in the dorsal region of the back,
for in this locality their places are usurped by stout
and tough ligaments which are quite inelastic. These
ligaments are much smaller and cord-like as they extend
between the low neural spines in the mid-cervical region.
Now it is only between the lofty neural spines of the
leading cervical vertebrse, commencing with the second
and third to include the fifth and fourth, and between
the last few cervicals that we see the true intersinnales ;
for passing the thirteenth cervical in ascending the
column, we find that these muscles gradually become
bilateral, and in the mid-region of the neck, stretch
between two vertebrae, and finally become continuous
with the anterior descending fasciculi of the longus
colli ])osticus.

136. The interarticulares represent another deep-
seated series of intervertebral muscles. Examining them
upon either side, we find that they start as a well-deve-
loped muscle extending between the postzygapophysis
of the axis to the hinder margin of the ring of the
atlas. Then in general throughout the vertebral column,
they occur as muscular bands extending from the
postzygapophysis of one vertebra to the same processes
of the vertebra next beyond. In old and muscular
subjects, however, we often see, l^etween the fifth

Tntertransver sails cervicis. Meckel, System, p. 294, No. 4.

Obliquus colli. Owen, Apteryx, p. 282.

Intertransversales anteriores et posteriores cervicis. Selenka, p. 99,

No. 17.
Transversus colli. Selenka, p. 94, No. 3.
Intertransverse muscles. Watson, p. 59."

Note. — In this connection see Owen (Anat. Verts., vol. ii., p. 89).
I question whether his obliquus colli represents the mm. intertrans-
versarii of Gadow.— R. W. S.

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294 THE MYOLOGY OF THE RAVEN.

and fourth, and fourth and third, and third and second,
that the fasciculi may extend so as to include two
vertel3r8e.

Owen is of the opinion that this series of muscles
in the Aj^teryx are the direct continuation of the
ohliquo-spinales of the back, in that bird.

137. The ohliquo-transversales are absent in the dorsal
region of the back in the Corvidce and our present
sul)ject. The Aptcryx possesses them, where they
" appear to be a continuation of the multijidus spincB
in the neck" (Owen).

In the Raven they are best studied as they connect
the ultimate cervical vertel^rse, say the ninth to the
twelfth inclusive. Here we find them passing obliquely
between the transverse process of one vertebra to the
postzygapophysis of the same side of the vertebra next
beyond but one.

They lie external to the short, straight inter-
articularcs, l^eing situated more on the sides of the
vertebrae, and are best brought into view l)y dividing
the main origin of the longus colli jiosticus, and care-
fully dissecting it away, as we proceed in the direction
of the head.

138. The intertransversales are represented by an
intimately connected though double series of short,
thick muscles, throughout the cervical division of the
colmnn, commencing between the third and fourth
vertebrae.

The stronger set extend between the transverse
processes of the contiguous vertebrae, wdiile the lower
or anterior set connect the consecutive parapophyses.

It wdll be seen upon dissection that this series of
muscles also fulfil another function, as they are so
arranged that they continue the lateral canals, or rather

THE MUSCULATUEE OF THE TRQNK. 295

afford tlie proper protection to the vessels and nerves
therein contained as they pass from one lateral canal
to the next, in the interspaces between the vertebrae,
where, of course, no osseous canal affords protection.

139. The triangularis sterni^ is a beautifully deve-
loped muscle within the cavity of the thorax.

It arises, somewhat tendinous, from the entire
superior margin of the summit of a costal process,
and its fibres directed backwards and upwards soon
divide into four well-defined digitations ; which latter
covering the inner surfaces of the leading three costal ribs,
become inserted into the first four as high up as their
articulations with the vertebral ribs.

It will at once be seen that, the sternum being fixed,
a contraction of these muscles will result in drawins:
down the costal ribs ; which, diminishing the cubical
contents of the thorax ; they thus become a j)owerful
auxiliary to the act of expiration.

The tiiangulai^is sterni in man, although it has
a little different origin, fulfils precisely the same
function.

^ According to Gadow, the following synonyms of the triangularis
sterni occur in literature {loo. cit., p. 125) : —

" 18/. M. COSTI-STERNALIS.

Der Erheber der Eippenfortsatze. Merrem.

Der innere oder dreieckige. Brustmuskel. (J/, sterno-costalis s.

triangularis sterni) Tiedemann, § 236.
Triangulaire du sternum. Cuvier.

„ „ „ Gervais et Alix, p. 16.

Sterno-costalis. Tiedemann.
Ohne Naraen. Meckel, System, p. 502.
Triangularis sterni s. sterno-costalis. Magnus, p. 226.
Sterno-costal. Alix, p, 386.
Triangularis sterni. Selenka, p. 104.

„ „ "Watson, p. 68."

296 THE MYOLOGY OF THE RAVEN.

140. Tlic intercostales^ are very prettily developed
ill a Raven. The fir.st or anterior one of the series
is the thickest and most evident. It arises from the
anterior free margin of the first dorsal ril3, commencing
as low down as its articulation with the costal rilj, and
is carried as high np as that point where the dorsal
rib gains its greatest width. The fibres pass obliquely
upwards and forwards, and are inserted into the pos-
terior margin of the last free rib, which is, of course, the

1 On pages 121, 122, and 123 of Bi-ona's Thier-Reichs (vi. Band),
Gadow devotes a very full account to two sets of " intercostal
muscles" in birds, his MiM. intercostales externi (18«) and Lis
Mm. intercostales intern: (18i), while in the same category he
places his Mm. iNXERArPENDicuLARES (18f). The synonymy of these
"rib-muscles" he presents as follows : — -

" h. Rippenmuskeln.

" 18. System der Mm. intercostales.

" Mm. intercostales.

Jh/sdes intercosfaux internes et externes. Vicq d'Azyr, pp. 292-293 ;

Cuvier, p. 324; Gervais et Alix, p. 13.
Mvi. intercostales externi et interni, o. innere und iiussere Zwischen-

I'ippenmuskeln. Tiedemann, § 239 ; Meckel, System^-^^. dO\

und 302, No. 1-5; Archiv, p. 248, No. 1-2; d'Alton, p. 13;

Owen.
Mm. intercostales externi et interni, o. innere und iiussere Zwischen-

rippenmuskeln. Magnus, p. 225 ; Selenka, p. 104, No. 2 ;

Gadow, No. 5 ; Watson, pp. 69 und TO.
Musculi interappendicidares costarum. Tiedemann.
Accessory external intercostal muscles. Watson, p. 70."

Under his (II.) system of the Yentri-laterale Muskeln (rt.
Halsregion), Prof. Gadow also describes his Mm. intertuberculares
(No. 17, p. 121): " Ein system von kurzen Muskeln als tiefste
Schicht der ventralen Halsmuskulatur. Sie beginnen als selbstiindige
Muskeln von den hinteren Halswirbeln an, wiihrend sie weiter
rlickwiirts innig mit den Mm. intertransversarii anteriores ver-
wachsen." And for a full account see the work refei-red to.

THE MUSCULATURE OF THE TRUNK. 297

one next in front of it. The remaining ril)s also have
intercostal muscles between them, but their fibres become
less and less better developed as we proceed back-
wards, and in all cases the strongest part of the muscle
is just above the articulations with the hsemapophyses ;
and from these parts the muscles also gradually grow
more feeble as we approach the vertebral column,
where they are su^^plemented by the levatores costarum.
Both an internal and external set can be made out.

Professor Owen, in his Anatomy of Vertebrates, does
not especially describe these muscles as' occurring in the
Apteryx.

141. The scalenus medius ^ is the most anterior leader
of that series of muscles which follow it, called the
levatores costarum, but is sufficiently conspicuous to
deserve a separate name and description. It arises from
the diapophysis and pleurapophysis of the eleventh
cervical vertebra, and its fibres passing downwards and
backwards are inserted upon the entire length of the short
free rib which articulates with the twelfth vertebra ; and
the middle fibres passing over and beyond it are inserted
into the anterior free margin of the middle third or more
of the second or longer free rib next behind it.

o

^ We have the subjoined synonymy of this muscle from Gadow
(loc. cit., p. 124) : —

" 18e. M. SCALENUS.

Les Tnuscles qui tiennent la jjlO'Ce des scalenes (pt.). Vicq d'Azyr.
M. scalene. Cuvier. {Als den Viigehi fehlend angefiilirt).
Scalenus (Rippenhalter). Wiedemann, p. 78.

„ (pt.). Tiedemann, § 223 ; Meckel, System, p. 301, No. 1 ;
Magnus, p. 220; Selenka, p. 98 ; Watson, p. 71.
Scalenus medius. Owen.
Surcostaux (pt.). Gervais et Alix, p. 12.

Musculus teres inter prima7n et secundum costam. Tiedemann, § 237.
M. teres. Magnus, p. 221."

298 THE MYOLOGY OF THE RAVEN.

This muscle is best developed, so far as I am at
present aware, in some of the Mammalia.

142. The levatores costanim^ constitute a series
of muscles following in sequence the scalenus raedius.
The first or anterior one arises from the extremity
of the transverse process of the twelfth vertebra, and
its fibres diverging as they pass downwards and back-
wards become inserted into the anterior free maroin of
the upper third of the long or posterior free rib, and
to the external surface of the same bone, contiguous to
this margin.

The succeedinor levator muscles of these ribs arise

O

and are inserted in a similar manner ; coming off" from
the ends of the diapophyses of all the dorsal vertebrae, and
being directed downwards and backwards are attached
to the anterior margins of the riljs next behind them,
in any case. They, however, grow gradually more and
more feebly developed as we proceed in the direction
of the pelvis, and are inserted less and less upon the
outer surface of the ri1j to the border of which they
are fast.

143. Tlie appendico-costales" represent anotl^er series
of thoracic muscles in birds, and are handsomely deve-
loped in the Eaven. In each case they arise from

^ Synonymy in Gadow is as follows [he. cif., j^p. 123, 124) : —

"18c?. Mm. levatores costarum.

Les vertehraux-costaux. Vicq d'Azyr.
Die Erheber der Biust. Merrem.
Levatores costarum. Tiedemann, § 235.

„ „ Owen.

,, „ Magnus, p. 222 ; Selenka, p. 98.

Heber der Rippen. Meckel, System, p. 301, No. 1.
Muscles sur-costaux. Gervais et Alix, p. 12."

2 These muscles are alluded to by Sir Richard Owen in his Anatomy

THE MUSCULATURE OF THE TEUNK. 299

the posterior edge of an epipleural appendage, and
forminof a thin sheet of muscle with its fibres directed
downwards and backwards, these latter become attached
to the outer surface of the rib next behind the one
bearing the appendage from which the muscle takes
origin. They are most powerfidly developed in the
anterior division of the series, growing less strong as we
proceed towards the pelvis, and of course, for obvious
reasons, the last pair of ribs do not possess them at all.
These epipleural appendages of the ri1)s are like-
wise supported, from above, by strong, triangular
aponeurotic membranes, which are attached to the
anterior edge of the process for its entire length, and

of Vertebrates, vol. ii., p. 92, while under his " Rippenmuskeln "
Gadow still describes another, viz. : —

" 18(7. M. QUADRATUS LUMBORUM.

M. quadratus lumhoruin. Tiedemann, § 241 ; Magnus ; Selenka,

p. 105, No. 31.
M. ohliquus abdominis internus (pt.). Gadow, No. 4.

" Der bei den meisten Eeptilien und Siiugethieren gewohnlich
stark entwickelte Quadratus lumborum ist bei den Yugeln sehr
reducirt. Er ist gewohnlich ein kleiner, diinner Muskel, der von
der Visceralfliiche des Hinterrandes der letzten falschen Eippe zum
Darmbeinkamme zicht. Er ist als die Fortsetzung der Intercostales
interni und des Obllquus abdom. int. in der lumbo-dorsalen Region
aufzufassen. Bisweilen sind dureh die Verkiimmerung der letzten
falschen Rippen zwei kleine Quadrati lumborum entstanden, z. B.
bei Rhea. Der eine mehr kopfwarts gelegene kommt vom Rande
des Tuberculum und dem proximalen Ende des Korpers der 2 Rippe
des 24 Wirbels und inserirt sich fleischig an den gleichen
gegeniiberliegenden Theilen der nachstfolgenden, fast ganz verkiim-
merten Rippe und an der Innenflache des Iliumkammes. Der
zweite Muskel entspringt von der Rippe des 25 Wirbels und inserirt
in Ermangelung eines Poippenbogens des 26 Wirbels fleischig auf
der Innenflache des Ilium in Hohe des 25 und 26 Wirbels " {he. cit.,
p. 126).

THE MUSCULATURE OF THE TEUNK. 301

are inserted into the posterior margin of the rib next
succeeding and anterior to it in any individual case.
With the first pair, these membranes are inserted on
the margins of the ribs nearly as high up as the
vertebral column, but grow shorter as we near the
pelvic extremity of the trunk,

144. The ohliquus externus abdominis^ is, as its name
implies, the most external of the layer of abdominal
muscles, and at the same time it is by far the most
extensive.

It arises by a delicate aponeurotic membrane from
the sides of all the true dorsal ribs, on a line drawn
at about the bases of their epij)leural appendages ; by
yri extension of the same membrane from the hinder
margin of the last vertebral rib, and the adjoining
margins of the pelvis ; by fleshy fil^res from the entire
posterior moiety of the inferior border of the post-
pubic element of the |)elvis ; and, fi.nally, again as-
suming the aponeurotic character, shades off" from the
structures lying about the root of the tail, and lower
part of the abdomen.

The aponeurotic membrane coming off from all

^ Most authors have described this muscle under this name, as
may be seen by the following synonymy (Gadow, loc. cit., pp. 126,
127):—

" 19. M. OBLIQUUS ABDOMINIS EXTEENUS,

M. oblique descende^is. Aldrovandi.

M. ohliquus exterior. Steno.

Le grand oblique. Yicq d'Azyr, p. 267, No. 1 ; Cuvier, p. 234 ;

Gervais et Alix, p. 17.
M. oblique descendens. Merrem, p. 151, No. 2.
M. ohliquus externus abdominis. Wiedemann, p. 79 ; Tiedemann,

§ 229; Owen, Apteryx, p. 286; Selenka, p. 102, No. 25;

Magnus, p. 230 ; Gadow, No. 1 ; Watson, p. 72.
Auesserer schiefer Bauchmuskel. Meckel, System, p. 303, No. 1 ;

Archiv, p. 249, No. 21."

302 THE MYOLOGY OF THE RAVEN.

the ribs, soon gives way at the lower thirds of the
ribs to a fleshy layer of transverse muscnlar fil3res
which are well developed. These in turn are also
more or less attached to the ribs beneath them, along
on their line of commencement below the epipleural
appendages. The muscular sheet thus formed passes
toward the sternum, loosely attached by fascia to the
hsemapophyses, overlying and concealing the latter
from sight as it does so. Again becoming aponeu-
rotic, it becomes attached to the side of the
sternum, and quite firmly so to the under side
of the lower part of the ]jectoralis major muscle
and the xiphoidal prolongation beneath it. The
transverse fibres of this muscular sheet extend down
the abdomen, and are continuous with those arising
from the edge of the posterior moiety of the post-
pubic element of the pelvis already alluded to above.
The muscular fijjres of the alidominal portion pass
somewhat downwards, though nearly transversel)^, to
the median abdominal line, where they terminate in
an aponeurosis just before arriving at the linea alba
For the upper two-thirds of the abdomen in the median
line, the aponeurotic fibres decussate, and are quite
firmly attached to the skin.

Owen says that in the Apteryx " the aponeurosis from
the last rib passes to be inserted into a strong ligament
extending between the free extremities of the pubic
bones, leaving the abdomen, behind the last rib, de-
fended only by the internal oblique and transver-
salis." As I have already said, in the Raven the very
delicate aponeurosis of the external ol)lique extends
into this recess between the last rib and the pelvis,
to become attached in the manner I have already
described.

THE MUSCULATUEE OF THE TRUNK. 303

145. The ohliqu'US internus cihdominis^ is exposed
after we have completely removed the external oblique.

It arises aponeurotic from the hinder third of the
post-pubic element of the pelvis ; and by fleshy
fibres from the remaining part of this bone ; and
occasionally by a few fibres from the iliac border beyond
the acetabulum. From this line of orio;in the fibres of
the internal oblique pass longitudinally over the abdo-
men to the thorax, where they become inserted into
the entire posterior margin of the last vertebral rib,
and into the pleurapophysial head of the last costal
rib ; which latter, as we know, is a floating one, and
articulates with this last vertebral rib.

The mesial margin of this muscle is free, and does
not blend with the rectus ahdominis, as Professor
Owen states that it does in the Kiwi-kiwi, nor do we
have the slightest clifliculty in the Eaven in dis-
tinguishing this muscle from the idtimate intercostal
muscle, as that eminent authority also found to be
the case in his subject.

146. The rectus ahdominis," on either side, arises

^ From the same source from which we obtained the synonymy of
the external oblique muscle of the abdomen, we have the following
(p. 127):-

" 20. M. OBLIQUUS ABDOMINIS INTERNUS.

Tertium par niusculortini abdominis. Aldrovandi.

M. ohliquus interior. Steno.

Le 2^6tit ohlique. Yicq d'Azyr, p. 267, No. 2; Cuvier, p. 324;
Gervais et Alix, p. 16 ; Alis, p. 387.

M. ohlique ascendens. Merrera, p. 151, No. 3.

M. ohUquus internus abdominis. Wiedemann, p. 80 ; Tiedemann,
§ 230 ; Owen, Apteryx, p. 286 ; Magnus, p. 231 ; Selenka,
p. 103, No. 26; Gadow, No. 4; Watson, p. 73.

Innerer schiefer Bauchmuskel. Meckel, System, p. 303, No. 2."

2 Following the synonymy of the muscles in Aves as given by

304 THE MYOLOGY OF THE RAVEN.

as an aponeurosis from the distal extremity of tlie
post-pubic element of the pelvis, and from that
semitendinous ligament which stretches from one
post-pubic ti]3 to the other, as far as the linea alba ;
the rectus of the opposite side arising from the re-
maining half of this ligament.

For half its distance, as the rectus ahdominis proceeds
towards the sternum, the muscle is purely tendinous ;
this tendon then terminates in nearly a straight
transverse line, from which the muscular fibres abruptly
commence. These latter then go, longitudinally,
directly to the xiphoidal margin of the sternum where
they are attached ; but beyond the lateral processes
of the sternum, though still in line with its sternal
attachment, the outer fibres of the rectus terminate
in a strong, aponeurotic membrane, which, as it passes
forwards is gradually lost over the outer surface of the
thoracic walls, beneath the external oblique and pectoralis
major muscles. The carneous portion of the 7'ectus
ahdominis in this bird fails to exhibit the slis^htest

Gadow we have the rectus cMominis thus compiled for us {loc. cit.,
p. 130 :—

" 23. M. RECTUS ABDOMINIS.

Par secundum inusculorura abdominis. Aldrovandi.
M. rectus (abdominis). Steno ; Mei-rem, p. 151, No. 1.

Wiedemann, p. 80.

Tiedemann, § 232.

Owen, Apteryx, p. 286.

Magnus, p. 232.

Selenka, p. 103, No. 27.

Gadow, No. 3.

Watson, p. 74.
Gerader Bauchmuskel. Meckel, System, p. 304, No. 4 ; Archiv,

p. 249, No. 23.
Le grand droit de I' abdomen. Gervais et Alix, p. 17.
„ „ „ „ „ Alix, p. 383."

THE MUSCULATUEE OF THE TRUNK. 305

evidences of the lincB transversce, so familiar to us in
many of the Mammalia ; and Owen also found them
to be present in the Kiwi-kiwi.

147. The transversalis abdominis'^ is a very well-
defined muscle in the Eaven, and is seen to arise,
thin and tendinous, from within the entire post-
pubic and iliac margins of the pelvis ; and to some
extent from the inter-pubic ligament. Becoming
gradually carneous, its fibres pass across the abdomen,
between the peritoneum, the rectus, and the internal
oblique, to become inserted over the entire pleural
aspect of the last two vertebral ribs, the intercostal
muscle between them, and the same surfaces of the
hsemapophyses connected below. This part of its
attachment is semitendinous. Its abdominal insertion
is into the linea alba, which just before reaching, the
transversalis becomes completely aponeurotic. This
aponeurotic area of the transversalis abdominis, when
taken in connection with the similar area of the muscle

^ That patriarch in anatomy, Aldrovandi, gave a striking name
to this muscle, as may be seen in the subjoined synonymy from
Gadow (loc. cit., p. 128) : —

"21. M. TRANSVERSUS ABDOMINIS.

Ultimum par quod transversaliuni in nobis locum obtinet. Aldrovandi.
Le muscle transverse. Vicq d'Azyr, p. 267 ; Cuvier, p. 324 ;

Gervais et Alix, p. 74.
M. transversalis. Merrem, p. 151, No. 4.
,, ,, Owen, Apteryx, p. 287.

„ ,, Watson, p. 74.

M. transversus abdominis. Wiedemann, p. 80.
,, ,, ,, Tiedemann, § 231.

„ ,, ,, Magnus, p. 232.

Selenka, p. 105, No. 30.
,, ,, ,, Gadow, No. 4.

Querer Bauchmuskel. Meckel, System, p. 304, No. 3 ; Archiv,
p. 249, No. 22."

X

306 THE MYOLOGY OF THE RAVEN.

of the opposite side, is of a spindle-like form, the
longitudinal axis of which is represented Ijy the
linea alba ; and its upper and lower apices being re-
spectively in the mid-point of the xiphoidal margin
of the sternum, and the mid-point of the inter-pubic
ligament. In a fresh specimen it is white and glistening,
and very characteristic.

148. The diaphragyn (Fig. 75) in the Kaven, as
in most existing birds, is in a rudimentary condition
only, and by no means meets the ends of that muscle
as we find it to be the case in Mammalia. It simply
consists of a thin, transparent membrane, overlying
the peritoneum on the one hand and the pericardium
on the other, completely conforming to the forms of
the various organs pressing upon it.

Three rudimentary muscles within the thorax, on
either side, play upon this attenuated midriff". These
arise from the vertebral heads of the second, third,
and fourth costal ribs, and their fil)res spread out
in a fan-like fashion upon the membrane in question.
These three muscles gradually increase in size as we
pass from before backwards, and are to be seen just
above the extremities of the digitations of the triangu-
laris sterni.

The best way to examine them is to disarticulate
the costal ribs of one side from the sternum ; care-
fully remove the muscular walls of the abdomen ; pull
the ribs away from the sternum, and pull down the
abdominal viscera, upon which the diaphragm will be
exposed, and the muscles in question at the sides of
the thoracic wall easily examined.

Macgillivray pointed out these muscles for us in his
figure showing certain anatomical structures as they
exist in Corvus frugilegus, and Coues has kindly re-

THE MUSCULATURE OF THE TRUNK. 307

produced this drawing for us in his Key to North
Aynerican Birds, the second edition (p. 206).

According to Sir Eichard Owen, " The diaj^hragm
presents more of its mammalian character in the Apteryx
than in any other known bird. It is perforated by
vessels only, in consequence of the non-development
of the abdominal air-cells. The origin corresponding
to that of the lesser muscles in Mammals is by two
strong and distinct, short tendinous pillars from the
sides of the body of the last costal vertebra ; they are
united by a strong tendon or fascia, forming the anterior
boundary of the aortic passage. The tendinous pillars
may be traced forward for some way in the central
aponeurosis, expanding without crossing ; they are
then lost in that aponeurosis, which is perforated by
the gastric arteries and veins, divides anteriorly to
give passage to the gullet and the apex of the heart,
expands over the anterior part of the thoracic air-cells,
and becomes, at its lateral circumference, the point of
attachment of muscular fil^res arising from the inner
surface of the anterior ribs, and forming apparently
a continuation of the transver sails abdominis " {A7iat.
of Verts., vol. ii. pp. 91, 92).

Up to the present time I have made no exhaustive
examinations of the dia/phragwi in other birds, as I hope
to on future occasions. For a number of reasons I
look forward to a study of its form in the Vultures
and Herons with no little interest.

149. The levator coccygis^ I believe to be the
detached caudal extremity of the longissimus dorsi

^ Seven muscles, as a rule, seem to control the movements of the
tail in most birds, I believe in all ordinary birds. These, as I have
already said in the text, I have attempted to give names, or choose
names for them from those they have already received at the hands

X 2

308 THE MYOLOGY OF THE RAVEN.

ill birds, which now in their present forms does
not extend beyond the postacetabular area of the

of others, such as would designate, not only their actions or
functions, but point out likewise their orderly arrangement as it
exists in the class for this part of the muscular system.

Gadow has given us quite a full synonymy of some of these
muscles, and by its aid no difficulty will be experienced in deter-
mining the corresponding muscles of the present work as I have
designated them. I give this synonymy below, with a few brief
remarks of its author under each heading (Bronn's Klussen des
Thier-Reiehs, vi. Band, pp. 131-135): — -

" d. Schwanzmuskeln.

" 24. M. LEVATOR COCCYGIS.

Par prinium ^musculorum uropygii. Steno.
Les deux releveurs du coccyx. Vicq d'Azyr, p. 274, No. 1.
Interepineux sacro-sus-caudien. Cuvier, p. 287, No. 1.
Grosser Schwanzheber. Merrem, p. 161, No. 62, No. 1 u. 2.
Levator coccyjis. Wiedemann, p. 82.

,, ,, Tiedemann, § 223.

Selenka, p. 100, No. 21.

,, ,, Gadow, No. 6.

,, ,, Watson, p. 65.

Levator cavdce s. spinalis caudoe. Gurlt, p. 19.
Schwanzheber und Zwischendornmuskeln. Meckel, System, p. 299 ;

Archiv, p. 247, No. 17.
Levator caudce. Owen, Apteryx, p. 286.

Sacro-coccygien superieur ; transversaire cpineux. Gervais et Alix,
p. 15.

" Bildet die dorsale Muskulatur des Schwanzes. Er entspringt bei
den Ratiten von der Spina iliaca, dabei manchmal weiter auf das
Os ilei und das Os ischii iibergreifend ; ferner von den Dornfortsiitzen
der ersten 3-4 Schwanzwirbel."

" Insertion an den Seitenflachen der Dorn- und an den Dorsal -
flachen der Querfortsatze der folgenden Schwanzwirbel. . . .

" 25. M. DEPRESSOR COCCYGIS.

Ahaisseurs du coccyx. Vicq d'Azyr, p. 274.

Niederzieher des Schwanzes. Merrem, p. 162.

Innerer Niederzieher des Steissbeines. Wiedemann, p. 82.

THE MUSCULATURE OF THE TRUNK. 309

dorsal aspect of the pelvis ; but, if we could liave
examined it in the now extinct avireptilian types, it

Depressor coccygis. Tiedemann, § 224 ; Selenka, p. 101, No. 24 ;

Gadow, Ratiten, p. 21 ; Watson, p. 67.
Ohne Namen. Meckel, System, p. \f\.
Depressor caudm. Gurlt, p. 19 ; Owen, Afieryx, p. 286.
Coccygien inferieur. Gervais et Alix, p. 16.

" Bildet die Muskulatur auf der Ventralseite der Schwanzvvirbel.
Entspringt fleischig von den Ventralfliichen der Querfortstitze der
letzten 8acral\virbel und der meisten folgenden freien Schwanz-
wirbel. Die einzelnen Blindel, die aber ahnlich wie die des M.
levator coccygis innig mit einander verwachsen konnen, icseriren
sich an den Ventralfliichen der nachstfolgenden Wirbelkorper. . . .

" 26 u. 27. Mm. pubi-coccygei.
" A. M. pubi-coccygeus externus + internus.
Moteurs lateraux chi coccyx. Vicq d'Azyr, p. 274, No. 2.
Oberer, grosser, und unterer ausdehnender Schwanzmuske].

Merrem, p. 162, No. 2 u. 5.
Puho + ischio-coccygiens. Cuvier, p. 287, No. 5 u. 6.
Sitzbein Schwanzmuskel. Meckel, Arcliiv, p. 248, No. 18 ; System,

§ 152.
Adductor caudcti inferior. Owen, Apteryx, p. 286.
Adductor caudce superior et inferior. Selenka, p. 131, No. 22.
Viscltio-p)tibo-coccygien. Gervais et Alix, p. 16.
Ischio-jmho-coccygeus. Watson, p. 67.

" B. M. puhi-coccygetis externus.
M. puhi-coccygetis. Tiedemann, § 226.

,, ,, ,, Wiedemann, p. 82.

,, ,, ,, Gurlt, p. 19.

M. pjuho-coccygeus externus. Gadow, No. 7.

" C. M. puhi-coccygeus internus.

M. ischio-coccygeus. Tiedemann, § 227 ; Gurlt, p. 19.
Depressor coccygis lateralis internus. Wiedemann, p. 82.
AI. puho-coccygeus internus. Gadow, No. 8.

" 26. M. PUBI-COCCYGEUS EXTERNUS.

" Dieser Muskel wird nur von der Haut und dem M, transverso-
analis bedeckt.

310 THE MYOLOGY OF THE UAVEN.

would have been found to be a continuation of the
last-named muscle.

" Er entspringt bei Rhea und Casuarius fleischig-sehnig von den
ventralen Flachen, der Enden der Querfortsiitze der ersten drei
Schwanzwirbel und dem distal-caudalen Ende der Ossa ischii et ilei.
Er geht dann quer liber den lateralen Theil des If. caud. il.Jiex.
fort, und in einem Bogen zum distalen Rande des Os pubis, woselbst
seine Fasern sicli mit denen des M. ohliquus et tranversus abdominis
vermischen. Bei Struthio ist er ganz mit dem M. jnibi-coccyy.
internus vereinigt.

" Die schwach entwickelte Steuerfedern besitzenden Carinaien
zeigen ein dem bei Rhea beschriebenen ahnliches Verhalten. Bei
den iibrigen Carinafen ist der Ursprung (oder Insertion) auf die
Ventralfliiche der Wurzeln der ausseren 2-4 Steuerfedern beschriinkt.

" 27. M. PUBI-COOCYGEUS INTERNUS.

" Innerster, eine breite und dlinne Schicht bildender Schwanz-
muskel auf der Seite des Baviches.

" Er entspringt in allegemeinen von der Ventralflache der
Querfortsiitze vmd der Korper der letzten freien Wirbel und der
Endplatte des Schwanzes. Er insei'irt sich, fjichei'formig ausgedehnt,
an der Innen- oder Ventralflache des distalen Theiles des Scham-
und benachbarten Sitzbeines, indem er sicli zwischen die distale
Grenze des M. obturator und des M. transversus abdominis schiebt.
Nacli Aussen wird er vom langen Kopfe des M. caud. il. fern, vom
M. jyubi-coccyg . extermis und vom M. transi'!erso-analis bedeckt,
wahreud er nach Innen dem Peritoneum autiiegt. . . .

" Bei LamelUrostres, Megalocephalon, Penelope ist er an den
Wurzeln der 4-5 inneren Steuerfedern befestigt, wiihrend nur
wenige sehnige Ziige zur Schwanzplatte gehen. Bei Picus viridis
fand ich ihu nur am ventralen Seitenrande der sebr stark ent-
wickelten Endplatte befestigt ; ahnlich bei vielen Singvogeln."

" 28. M. ILIO-COCCYGEUS.

M. ileo coccygien. Cuvier, p. 287, No. 4.
Quadratus coccygis. Selenka, p. 101, No. 23.

" Zum System der ventralen Schwanzmuskulatur gelicirt auch der
M. ilio-coccygeus, da er wie der M. p)ubi-coccygeus durch Aeste aus
dem Plexus pudendus innervirt wird, trotz seiner dorsalen Lage.

" Er erscheint nach Wegnahme der Haut auf der dorsalen Seite

THE MUSCULATURE OF THE TRUNK. 311

In the RavGD, it arises, on either side, from a limited
area of the ilium just beyond and to the side of the an-
terior free caudal vertebra. The fibres converge, and end
in a tendon which becomes inserted into the tul)erosity
on the anterior margin of the pygostyle. In passing the
neural spines of the other caudal vertebrae, however, it
throws off to each one a tendon which is inserted upon
their several apices, though somewhat hidden from
view by the muscle itself.

It depends upon the muscularity and age of the
subject, as to how far forwards this muscle encroaches
upon the pelvis for its origin ; the older and more mus-
cular the subject, the more extensive the encroachment
and area of origin, as a rule. As already stated, this
muscle is continuous from back to tail in Reptiles
generally.

150. The levator caudce is a lono- oblono; muscle,
considerably larger than the last, and lying immediately
to its outer side, and cpiite intimately connected with
it for its entire length by a firm fascia.

It, too, arises from the hinder surface of the post-
acetabular area of the pelvis — but not as far forwards
as the levator coccygis — as well as from the superior

des Schwanzes, seitlich neben dem M. levator caudce. Er entspringt
fleischig von der dorso-medialen Flache des distalen Ilium, und von
der Dorsalflache der Querfortsatze der meisten Schwanzwirbel. Er
inserirt sich an der Dorsalflache der 4-5 aussersten Steuerfedern,
welche er spreizt und hebt.

" Bei Picus viridis setzt er sich nur an die beiden aussersten
Steuerfedern.

" Der M. ilio-coccygeus und die Mm. pubi-coccygei gehbren eigent-
lich nicht zu den echten (spinalen) Schwanzmuskeln, da sie am
Becken inseriren und aus dem Plexus pudeTidus innervirt werden.
Sie bilden daher den Uebergang von den Muskeln des Stammes
zu denen der Extremitaten-Giirtel."

312 THE MYOLOGY OF THE RAVEN.

surfaces of all the vertebrae of the tail, except the
pygostyle. It is inserted into the quill-butts of the four
rectrices that lie next in order to the jDygostyle, which
insertion gives it the power of forcibly elevating these
four feathers by its contraction.

151. Tlie transversus perinei^ arises, on either side,
from the entire posterior margin of the ischium, and
from the posterior margin of tliat portion of the post-
pubis which extends 1 )eyond it. This somewhat tendinous
origin makes a loop at the salient angle where the
ischium and ilium unite behind, through which loop
the femoro-caudal muscle passes. From this point of
attachment a dense fascia extends towards the coccyx,
being closely attached to the entire hinder margin of
the ilium, and firmly holding down the lower strata
of the coccygeal mucles. From this line of origin the
fibres composing the thin and sheet-like transversus
'pervnei converge and pass towards the median line,
to unite in a raphe with the muscle coming from the
opposite side, in front of the anus.

These muscles serve to support the viscera here at
the lower part of the abdomen, and so, too, perform
the function of that muscle found in certain of the
Mammalia know^n as the levator aiii.

152. The depressor caudcB is a strong conical muscle

^ But few comparative morphologists seem to have designated
this muscle by the name I here bestow upon it, although Owen, I
believe, used it. Gadow {loc. cif., p. 129), presents us Avith the
following brief synonymy only, viz. : —

" 22. M. TRANSVERSO-ANALIS.

M. levator ani ; M. transversus ossium pubis. Gurlt, p. 20.
M. transverso-cloacalis. Gervais et Alix, p. 16.
„ ,, „ Watson, p. 71.

Aufheber des Afters. Tiedemann, § 430."

THE MUSCULATURE OF THE TRUNK.

313

wliicli overlies the depressor coccygis, and one that
is devoted to pulling the tail downwards and out-
wards, and by an equal contraction of the muscle of
the opposite side, directly downwards.

cL-icLp7u^cLgrrv.

Fig. 75. — An anterior portion of the trunk of a Raven, with the sternum, muscles,
and other parts extensively cut away in order to show the rudimentary
muscles of the diaphragm. The heart is pushed somewhat forward, and
tensely covered by its membranes, &c. The dissecting-chains pull the
lateral chest- walls apart. Life-size, by the author, from his own
dissections.

It arises from the lower half of the posterior border
of the ischium, and from the entire posterior border of

314 THE MYOLOGY OF THE RAVEN.

the post-pubis beyond it, all of wliicli is beneath the
origin of the transversus perinei. The fibres converge
as they pass backwards and inwards, and turn in such
a way as to permit the muscle to become attached to
the quill-butts of the three or four outer rectrices.

This muscle must be cut across, and its extremities
reflected, before we can thoroughly examine the next
one.

153. The depressoi' coccygis arises from rather more
than the lower half of the posterior margin of the ischium,
as well as from the anterior three-fourths of the posterior
margin of the post-pubic element of the pelvis imme-
diately beyond it. This line of origin lies well within
the marginal border of the bones mentioned, as the pre-
ceding muscle is more properly attached to their free
edges.

Its fibres converge as they pass backwards and
inwards, and are inserted, somewhat tendinous, to the
thickened rim of the inferior and expanded portion
of the pygostyle, on the side corresponding with
the origin of the muscle.

154. The lateralis caudcB is composed of usually four
fasciculi, more or less joined together by their inter-
muscular fasciae, the external fasciculus being by far
the most distinct one. This arises from the tip of the
transverse process of the first free caudal vertebra at
the anterior end of the series, and is inserted on the
outer side of the proximal extremity of the external
rectrix of the tail. The succeeding fasciculi of the
lateralis caudcB arise, in order, from the tips of the
next three transverse processes of the caudal vertebrae
following the one just alluded to. Their fibres pass
backwards, and are inserted on the under side of the
quill-butts of the first two or three outer rectrices.

THE MUSCULATURE OF THE TRUNK. 315.

The combined muscle forms a fleshy mass, at the
side of the skeleton of the tail ; and it is evident that
when the outer fasciculus of the muscle alone contracts,
as it may, it will, assisted by an equal force exerted
on the part of the corresponding fasciculus of the
muscle of the opposite side, tend to powerfully pull the
tail feathers apart, and thus spread them. To a certain
extent the remaining fasciculi can act in the same
way, but they are inserted in such a manner that
by their contraction these rectrices will he pulled both
downwards and outwards.

155. The lateralis coccygis is a powerfully developed
muscle found immediately beneath the ends of the
transverse processes of the caudal vertebrae, and where
it is attached to the expanded part of the pygostyle
it blends more or less with the muscle of the opposite
side, and the two in this region are firmly braced
down by a confining, white and glistening, tendinous
fascia, which is at once made conspicuous by the
removal of several of the muscles described in the fore-
going paragraphs.

The lateralis coccygis arises from the nether aspect
of the posterior end of the ilium, and by tendons
which severally spring from the under side of the
ends of the leading three, occasionally four, caudal ver-
tebrae. The combined fibres form a subcompressed
muscular mass, directed backwards and slightly inwards,
to become attached, on either side, to the posterior mar-
gin, exclusive of the midspine, of the expanded portion
of the pygostyle beneath.

This pair of muscles control the lateral movements
of the tail and the feathers attached to it, and to
some extent its oblique downward movements.

156. Tlie infracoccygis is that muscle which is

*

31G THE MYOLOGY OF THE RAVEN.

found l)cneatli the skeleton of the tail, and which
beyond the fifth caudal vertebra mc^rges to a great
extent with the fellow of the opposite side, almost
giving it the appearance of an azygos muscle.

It arises, by means of a strong tendinous at-
tachment, on either side, from the inferior surface
of the diapophysis of that last vertebra which
anchyloses with the pelvic sacrum. It also finds
origin from similar positions on the first four free
caudal vertebree ; this part of the muscle seems to
merge with the one of the opposite side to form a
tendinous attachment to the prominent hypapophysis
of the fifth caudal vertebra. But notwithstanding
this, the same muscle seems to spring again from
the under side of the transverse processes of the
remaining caudal vertebrae, and here is where the
blending seems to take place, not only with the oppo-
site muscle, l)ut as a continuation of the first part
described. Finally, this latter fleshy portion becomes
inserted into the lower sides of the pygostyle, and
by a tendon on its middle infero-spine, or aborted
hypapophysis. ,

This pair of muscles evidently antagonize the leva-
tors found on the u^iper side of the caudal vertebrge
and sacrum.

Sir Kichard Owen, in his description of the caudal
muscles as they were found by him to exist in the
Apteryx, bestowed names upon them quite different
from those that I propose for them here. In the
nomenclature of this anatomist we find such names
as the ischio-coccygetis, the quadratics coccygis, the
puho-coccygeus, and others. A careful study on my
part of the musculature of this part of the skeleton and
its rectricial appendage in the Raven, convinced me that

THE MUSCULATURE OF THE TRUNK.

317

a far better nomenclature might be adoped for these
muscles, as, notwithstanding their various modes of origin
and insertion, they were, in this bird at least, evidently
arranged upon quite a definite plan.

Of the two pair of levators one is devoted to the
skeleton and one to the rectrices, and these are directly

Fig. 76.— Under view of posterior portion of pelvis of a Raven, together with the
caudal vertebrae ; designed to show the position of the infracoccygis
and lateralis coccygis muscles. Life-size, and drawn by the author
from his dissections.

antagonized by the pair each of which I have called an
infracoccygis.

Now we still have remaining lateral muscles both
for the skeleton of the tail and the rectrices, while to
these are added depressors for the same parts, and
omitting the transversus jyerinei, the whole is com-
pleted by the femoro-caudals, which lead from the
pygostyle to a thigh-bone on either side.

318 THE MYOLOGY OF THE RAVEN.

A moment's reflection is sufficient to convince us that
all those varied movements, so essential to the feathery
rudder of this black avian knight, can be executed to
a nicety by this wonderfully complete musculature of
his caudal extremity.

BIBLIOGRAPHY.

Important works and papers treating of the muscles of birds,
compiled, abridged, and rearranged from the bibliographical lists
of Hans Gadow, and several other sources, as well as many new
titles added thereto by the present writer.

1. Aldrovandi, Ulysses, Be musctdis aquilce: Or7iithologia, tom. i.,

p. 117. Bononite, 1599, folio; Frankfurt, 1610.

2. Alix, E., Sur le membre abdominal des oiseaux. Bullet.

de la SoG. pldlomat., 1864.

3. Comparaison des os et des muscles des oiseaux avec ceux

des mammiferes. Op. cit., 1867.

4. Muscles flechisseurs des orteils chez les oiseau. Op.

cit., 1874.

5. Sur la determination du muscle long supinateur chez les

oiseaux. Jour, de Zoologie, iii. (1874), pp. 21-25.

6. Essai sur Vajypareil locotnoteur des oiseaux. Paris, 1874.

Planches i.-iii.

7. Memoire sur I'osteologie et la myologie du Nothura

major. Jour, de Zoologie, iii., 1874, pp. 167-214, 252-
282, pis. viii.-xi.

8. Sur quelques points de I'anatomie du Nandou. Ibid.

9. Sur la myologie du Rhynchotus rufescens. Jour, de

Zoologie, v. (1876), p. 411.

10. d'Alton, E., Be strigum musculis commentatio. 4to. Halis,

1837.

11. ^i.k?,\\]^, G., Anatome Animaliaim. Amstelodami, 1681.

12. BoRELLi, G. A., Be motu animalium. Romse, 1680.

13. Burmeister, H., Systematische Uebersicht der Thiere Brasiliena,

2 Theil, i. Halfte. Berlin, 1856.

320 BIBLIOGRAPHY.

14. Burton, E., Observations on the Natural History and Anatomy

of the Pehcanus {Tachypetes) aqulla. Trans. Lian. Soc.
Lond., vol. xiii., pp. 1-11 (1822).

15. Carlsson, Albertina, Beitriige zur Kenntniss der Anatomie

der Schwimmvcigel, 5 Taf. K. Svenska Vet. Akad.
Handlincjar, Bd. 9, ISTo. 3, 1884.

16. Carus, C. G., Erlduteru)igstafeln zur vergleicltenden Anatoiide,

Heft i. Leipzig, 1826. Taf. iv.-v.

17. CouES, E., Osteology of Colymhus torquatus, with Notes on its

Myology. Mem. Boston iSoc. of Nat. Hist., i. (1867),
pp. 131-172.

18. Cunningham, E. O., Notes on some points in the Anatomy

of Three Kingfishers. P. Z. 8. Lond., 1870, pp. 280-283,
p. xxiv.

19. CuviER, G., Le<ions d' anatomie comparee, 3 edit., tome i. Paris,

1835.

20. DoNiTZ, W., Ueber die Halswirbelsiiule der Vijgel aus der

Gattung Plotus. Arcldvf. Anat. und Pliysiol., 1873, pp.
357-360, pi. ix.

21. DuvERNOY, G. L., Memoires sur quelques particularites des

organs de la deglutition de la classe des oiseaux et des
reptiles. Mem. de la Societe d Histoire naturelle de Stras-
bourg, tome ii., Paris, 1835, with 5 plates.

22. Kiirzere Notizen in Comptes rendus de V Academic des

Sciences, tome ii., 1836, pp. 187-191.

23. Edwards, A. Milne-, KecJierches anatomiques et paleontologiques

pour servir cb Vhistoire des oiseaux fossiles de la France,
tome i., planches ix.-x. Paris, 1867-68.

24. Edwards, H. Milne-, Le<^ons sur la physiologie et V anatomie com-

paree deVhomme et des animaux, tome xi. Paris, 1875.

25. Forbes, W. A., Collected Scientific Papers. London, 1885.

26. FiJRBRiNGER, M., Zur Lehre von den Umbildungen der

Nervenplexus. Morpholog. Jahrhuch, Bd. v., 1879,

p. 358.
27. Untersuchungen zur Morphologic und Systematik der

Vogel, zugleiclh ein Beilrag zur Anatomie der Stiitz- und

Bewegungsorgane. Amsterdam, 1888.
28. ■ Ueber das Schulter- und EUenbogengelenk bei Vbgeln

und Reptilien. Morph. Jahrh., xi., 1885, pp. 119-121.
29. Ueber Deutung und Nomenclatur der Muskulatur des

VogelflUgels. Morph. Jahrh., xi., pp. 122-126.

BIBLIOGRAPHY. 321

30, FiJRBRiNGER, M., MonograpMe der Schtdter uml der Flug-
'inuskeln der Vogel. 4to. 1885. Plates iv.

3J. Gadow, H., Zur vergleichenden Anatomie der Musladatur des
Beckens und der hirderen Gliedmasse der Ratiten. 1880.
5Taf.

32. Untersuchungen liber die Bauchmuskeln der Kro-

kodile, Eidechsen und ScliildkriJten. Morphol. Jahrhuch,
vii., 1882, pp. 57-100, Taf. vi.

33. Beitriige zur Myologie des Beckens und der hinteren

Extremitat der Reptilien. Morphol. Jahrhuch, vii.,

1882, pp. 329-466, Taf. xvii.-xxi.

34. On some points in the Anatomy of Pterodes arenarhis,

with remarks on its systematic position. P.Z.S., 1882,
pp. 312-332.

35. Observations in Comparative Myology. Jour, of Anat.

and Physiology, 1882, pp. 493-514.

36. Vogel : Aves. Bronn's Klass. u. Ord. des Thier-

Reichs, vi. Band, iv. Abtheil. Leipzig and Heidelberg,
1888.

37. On the Suctorial Apparatus of the Tenuirostres. P.Z.S.,

1883, pp. 62-69, pi. xvi.

38. Garrod, a. H., The Collected Scientific Papers of. London,

1881.

39. Geoffroy Saint-Hilaire, L, Philosopjhie anatomique, tome i.

Paris, 1818.

40. Gervais, p., et Alix, E., Osteologie et Myologie des Manchots.

Journal de Zoologie, tome vi., 1877.

41. GiEBEL, C., Bemerkungen iiber Cafhartes aura, Falco albi-

cilla, F. lagopus, und F. huteo. Zeitschr. f d. gesammt.
N'aturio.,Bd. ix., 1857.

42. Zur Naturgesehichte des Surinamischen Wasserhuhnes

(Podoa surinamensis). Op. cit., Bd. xviii., 1861, p. 424.

43. Ueber einige Nebenknochen aus Vogelskelett. Op. cit..

Bd. xxviii., 1866, p. 29.
44. • Die Zunge der Vogel und ihr Geriist. Zeitschrift

gesammt. Naturwiss., Bd. xi. (1858), pp. 19-51,

Taf. i-viii.

45. Gruber, W., Beohacldungen zur menscldichen und vergleichenden

Anatomie, ILeit 2. Berlin, 1879, p. 45.

46. GuRLT, E. F., Anatomie der Ilausvbgel. Berlin, 1849, 8vo,

Taf. i.-v.

Y

322 BIBLIOGRAPHY.

47. Hartwig, p., Observations sur rentendue relative des ailes

et le poids des muscles pectoraux chez les animaux
vertebres volants. Archive neerlandais des Sciences
exactes et natureUes., tome v., La Haye, 1869, p. 31.

48. Haswell, W. a., Note on some Points in the Anatomy of the

Pigeons referred to by Dr. Hans Gadow in a recent
paper on the Anatomy of Pterocles. Proc. Linn. Soc.
of New Soutli Wales, vol. vii., p. 3, p. 397-402.

49. Some Points in the Myology of the Common Pigeon.

Jour. Anat. and Physiol., vol. xvii., p. 11 ; Jan., 1883,
pp. 218-221, p. Ill ; April, 1883, p. 404.

50. Haughton, S., Notes on Animal Mechanism : No. 3, On the

Muscular Anatomy of the Leg of the Ostrich. Proc.
Roy. Irish Acad., ix., 1865, pp . 50-61 ; also in Ann.
3£a(j. Nat. Hist., 3 ser., xv., 1865, pp. 262-272.

51. No. 10, Muscular Anatomy of the Emu. Proc. Roy.

Irish Acad., 1868, pp. 487-497.

52. No. 11, Muscular Anatomy of the Rhea. Ibid., pp.

497-504.

53. No. 17, On the Comparative Myology of certain Birds.

Ibid., pp. 524-526.

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INDEX.

INDEX.

All the muscles described in this work are indexed in alphabetical order under
the word "Muscle," and those that have not received a special description are, as a
rule, indexed under those letters that begin their respective naynes. Usually they
are muscles not occurring in the Birds which have been myologieally treated in the
present volume, and often have a nmnenclature not generally in use.

Under the words "Synonymy of" have been indexed in alphabetical order the
majority of those muscles whose synonyms have been given in footnotes throughotot
the work.

A, SYMBOL for femoro-caudal, 188
Abductor indicis, of Gadow, 149
Abductor longiis hallucis muscle of

Apes, 214
Accessory metatarsal bone, 246
Action of entotympanic muscle, on

raising superior mandible, 19
obturator muscles in Kaven,

194

pectoralis major muscle, 70

pectoralis secundus muscle in

Raven, 73
pronator muscles of forearm,

132
Agouti, soleus muscle in, 208

tibialis anticus in, 214

Air passages, muscles of, 43
Albatrosses, pectoralis secundus

muscle in, 74
Alcedinidoe, 273
Aldrovandi, work cited, 319
Alix, work cited, 319
Alton, d', work cited, 319
Anliinga, nuclial style in, 265
Annular ligament of foot, 230
Annularis, term defined, 241
Anolis, caudal muscles in, 261

Anseres lack accessory semitendi-

nosus, 181
Anserine fowl, air passages in, 52
Anterior fasciculus of gluteus

primus, 159
Anthropoid Apes, extensor ossis meta-

carpi pollicis muscle in, 133
— — tibialis anticus muscle in, 214
Apes and Man, separation of flexors

of forearm in, 141

pectoralis minor muscle in, 100

Aponeurosis, formed by certain

muscles of thigh in Raven,

158
Apteria in Raven, 2
Apteryx, 13

abdominal muscles in, 302

biventer cervicis in, 271

certain muscles in (described

by Owen) not identified,

197, 198

dermal muscles in, 8, 9, 13

— — ■ diaphragm in, 307

flexor metarcarpi ulnaris in,

144

gemellus muscle in, 194

gluteal muscles in, 166

interarticularis in, 294

longus colli in, 287, 288

332

INDEX.

Apteryx, longus colli posticus in,

275

mantelli, 239, 240

• obliquo-transversalis in, 294

obliquus colli in, 284

Owen's description of dorsal

muscles in, 284, 285

■ pectoralis major muscle in, 71

peroneus longus muscle in,

210, 211
■ rhomboideus muscle absent in,

85

• sacro-lumbalis in, 278

semitendinosus muscle in, 179

serratusmagnusanticus muscle

in, 99

soleus muscle in, 206

subscapularis muscle in, 87, 104

■ teres et infraspinatus muscle

in, 91

■ tracihelo-mastoideus in, 290

trunk muscles in, 259

Archteopteryx, 261

Area of obturator internus, form of,

as used in taxonomy of

birds, 191, 193
Arteries, gastric, in Apteryx, 307
Auks, bicipital slip to patagium in,

109

B

B, SYMBOL for accessory femoro-

caudal, 188
Basal joint of hallux in Raven, 201
Batracliians, ventral muscles in, 262
Bats, trapezius muscle in, 108
Bell, Jeffrey, work cited, 63
Bicipital slip to the patagium, 109
Birds, tibialis anticus in, 214
Blasius, work cited, 319
Bones, arytenoid, 45
Borelli, work cited, 319
Bronchial half-rings in Raven, 43
Burmeister, work cited, 319
Bursa over trochanter of femur, for

tendon of gluteus medius,

163, 169
Burton, work cited, 320

Carlsson, work cited, 320
Carotid artery in Raven, 268
Carus, work cited, 320
Cassowary, its gluteal muscles, 167
Caudal muscles in Raven, 261

recapitulation of, 317

Centre of gravity of bird's body, 73
Ceryle, biventer cervicis in, 273
(Jetacea, 262
Cetaceans, subscapularis muscle in,

104
Chamasleon parsonii, 261
Chameleo, tibialis anticus in, 214
Chameleon, 276
extensor ossis metacarpi pol-

licis muscle in, 133
Chauna, expansor secundariorum

muscle in, 78
Chelonians, muscles of the dorsum,

259
Ciconine character, expansor secund-
ariorum muscle, 78
Claus, work cited, 63
Colymbus septentrional is, 264
Coraco-clavicular membrane, 76, 77
Cormorant, osseous style of cranium

in, 18
Cormorants, nuchal style in, 265
Coracoid, kept in place by sub-

clavius muscle, 95
Corvidte, bicipital slip to patagium,

absent in, 109
Corvus frugilegus, diaphragm in, 306
Coues on muscles and tendons in

legs of birds, 225

works cited, 306, 307, 320

Cranes, air passages in, 52

bicipital slip to patagium in,

109
Cricoid bone in Raven, 45
Crocodile, depressor palpebroe in-

ferioris muscle in, 56
Cunningham, Dr., works cited, 273,

320
Cuvier, on sterno-trachealis muscle,

49

■ work cited, 320

Cyanocephalus cyanocephalus, ob-
turator muscle in, 192

C

Caprimulgi, bicipital slip to pata-
gium in, 109
ej'e closed by upper lids in, 57

D

Deglutition, act of, as affected by
action of stei'no-hyoideus
muscle, 30

INDEX.

333

Dermal muscles, method of exposing

for dissection, 3

number of, 3

■ ■ two varieties of, 2

Diaphragm, muscles of, in Raven,

313
Dissection of muscles of lower ex-
tremity in Raven, 155
of tendons in foot of Raven,

method of, 222, 223, 225
Diurnal birds of prey, pectoral

muscles in, 71
Donit/, work cited, 320
Dorsal muscles of upper extremity of

Raven, 78
Drum of ear in Raven, ligament

attached to, 62
Duckbill, 259
Ducks, expansor secundariorum

muscle in, 79
Duvernoy, work cited, 320

E

Eagles, semitendinosus muscle ab-
sent in, 181
Ear- conch, ligaments of, 5
Ear, muscles of, 53, 62
Echidna, 259

■ gemellus muscle absent in, 194

subscapularis muscle in, 104

tibialis anticus in, 214

Edwards, Milne-, peronei muscles of,

228

works cited, 320

Elbow-joint, 12, 13

Emyp, 277

Epipleural appendages, 13

Eustachian tube, relation of tensor

tympani to, in an Owl, 63
Extensor brevis digitorum of Owen,

219, 220
Extensor longus digitorum, separate

slip of, 217
Extensor plicae alaris of Owen, 106,

125
Extensor pollicis brevis, 220
Extensor proprius hallucis in man,

148

of Mivart, 220

External malleolus, 228

Eye, method of dissection of muscles

of, in Raven, 53
muscles of, 53

F

Fibrous sheath for muscles, distal

end of ulna, 129
Fieldfare, Owen's figure of tongue-
muscles in, 33
Fishes, subvertebral caudal muscles

in, 262
Flexors in leg of Raven, how named,

227
Flexor metacarpi ulnaris, of Owen,

144
Flexor minimi digiti brevis, as a

posterior ligament to little

finger in Raven, 150
Flexor perforatus annularis secundus

pedis, 252
Flexor pollicis of Gadow, 150
Flying Squirrel, modification of

platysma myoides muscle

in, i08
Foramen for sixth nerve in Raven,

59
Forbes on muscles and tendons in

legs of birds, 225, 226
on semitendinosus muscle in

birds, 179

on the ambiens muscle, 171

work cited, 320

Formulae for muscles of thigh in

birds, as used in classifica-
tion, 188
Fowl, vinculum in, 240
Frog, tibialis anticus in, 214
Function of subclavius muscle in

Raven, 94, 95
Fiirbringer, on the dermal muscles, 3

works cited, 320

Furnaria, sterno-trachealis muscle

in, 51

G

Gadow, on muscles of eye and ear, 63

on the dermal muscles, 3

on the digastric and pterygoidal

muscles, 15, 16

works cited, 321

Galeopithecus, modification of pla-
tysma myoides muscle in,
108

Gallinaceous birds, air passages in,
52

pectoralis tertius muscle in, 76

334

INDEX.

Gallinse, expansor secundarioruni

muscle in, 79

longus colli posticus in, 275

Gallus, 238, 239

Garrod, on muscles and tendons in

legs of birds, 225, 226

work cited, 234, 235, 321

Geese, expansor secundarioruni

muscle in, 79
Gegenbaur, work cited, 63
Geococcyx californianus, accessory

semitendinosus in, 182

biceps flexor cruris in, 175

extensor brevis digitorum in,

220
extensor fenioris in, 175, 176,

177
extensor longus digitorum in,

217
flexor muscles of the leg in,

251
flexor perforatus indicis se-

cundus pedis and flexor

longus hallucis muscles in,

233, 234
gastrocnemius muscle in, 203,

204
gluteal muscles in, 167, 168,

169
obturator, gemellus, and ad-
ductor muscles in, 198, 199
■ peroneus longus and tibialis

anticus muscles in, 212, 213,

214

popliteus in, 250

sartorius muscle in, 157

semimembranosus muscle in,

182, 183
— — - semitendinosus muscle in, 181

soleus muscle in, 206

tibialis posticus in, 229

Geoffroy Saint-Hilaire, work cited,

321
Gervais, work cited, 321
Giebel, work cited, 321
Gigantic Crane, upper larynx in, 47
Gluteus maximus, of Cuvier, 158

of Meckel, 158

Gluta3us quartus, of Mayer, 167
Glutpeus cjuintus, of Mayer, 167
Goose, coraco-brachialis muscle in,

96

longus colli posticus in, 275

■ mesial aspect of eye in, 55

pyramidalis nictitantis in, 57

Gracilis muscle, 171

Greater 6-igmoid cavity of ulna, 133

Grebes, semimembranosus muscle

in, 181
Gruber, work cited, 321
Guillemots, pectoralis secundus

muscle in, 73, 74
Gull, Selenka finds soleus muscle in,

205
Gulls, bicipital slip to patagium in,

109
pectoralis secundus muscle in,

73, 74

pectoralis tertius muscle in, 76

Gurlt, work cited, 321

H

Hartwig, work cited, 322

Haswell, work cited, 322

Haughton, work cited, 322

Heart of Eaven, 313

Heming, work cited, 322

Herissant, work cited, 322

Heron, coraco-bracliialis muscle in,
96

Herons, diaphragm in, 307

jiectoralis major muscle in, 71

Heusinger, work cited, 322

Homo, tifth layer of muscles of dor-
sum in, 284

flexor minimi digiti muscle

in, 148

muscles of back in, 276

Hoopoe, vinculum absent in, 235

Horse, extensor ossis metarcarpi pol-
licis muscle in, 134

Huber, work cited, 322

Humboldt, work cited, 322

Humero-ulnar pulley, 142

Humerus, 13, 14

various muscles attached to, in

Raven, 83

Humming-birds, not especially re-
lated to the Swifts, 189"

pectoralis major muscle in, 71

Humphrey, work cited, 322

Hunter, work cited, 322

Huxley, work cited, 63. 322

Hyaena, soleus muscle absent in,
208

Hyoid apparatus, 22, 23

Hypotarsus, 209

Hyrax, tibialis anticus in, 214

INDEX.

335

Icterus vulgaris, patagial muscles

in, in, 113
Iguana, 276
Iguana tuberculata, 261
Ilio-neural canal, 279
Ilium, 12

Intermetacarpal fenestra, 154
Interorbital vacuity, 56
Interosseous membrane of forearm,

128, 133
Interpleurapophysial membrane, 93
Instruments used in dissection of

birds for the muscles, 1
Intestinal ca3ca absent in Trocliili

and Cypseli, 189
Iscliiatic foramen, and obturator

externus muscle, 186

Jacquemin, work cited, 322

Jaeger, work cited, 323

Jaw, muscles that act upon, 19

ramus of, origin of platysma

myoides. 6

K

Kaczander, work cited, 323

Keel of sternum, muscles attached

to, 70, 72
Kiwi- kiwi, levator scapulae muscle

in, 99

linae transversa; in, 305

obliquus internus abdominis

in, 303
Klein, work cited, 323
Klemm, work cited, 323
Knee-joint, fascia of, 157
Kutorga, work cited, 323

Lacrymal bone, origin of orbicularis
palpebrarum on, in Raven, 54
Lanth, work cited, 323
Larus fuscus, soleus muscle in, 205
Legal and Eeichel, work cited, 323
Ligamentum nuchse, 265
Ligamentum patellae, 157

LinjB trans versoe, absent in Raven,

305
Linea alba, 302, 305, 306
Linea aspera, 184, 199
Lingual apparatus, 6, 26
Liolepis belli, 261
Lower extremity, muscles of, 155,

199
Lower larynx in Raven, 44, 47, 50
method of examining, in Raven,

47
Ludwig, work cited, 323

M

Macalister, work cited, 323
Macgillivray, on diaphragm in

birds, 306
Magnus, work cited, 323
Mammalia, diaphragm in, 306

eye-muscles in, 58

levator ani muscle in, 312

• ligamentum nuchoe in, 265

longus colli anterior in, 286

mylo-hyoideus muscle in, 26

Mammals, peronei muscles in, 226

serratus magnus muscle, 105

subscapularis muscle in, 104

Man, de, work cited, 323
Man, flexor profundus digitorum
in, 141

■ ■ tibialis anticus in, 215

Mandible, muscles attached to, 21
Mayer, Professor, on gluteal muscles

of the Cassowary, 166, 167
Mayer, work cited, 323
Maxillary bone, origin of orbicu-
laris palpebrarum on, in
Raven, 54
Meatus auditorius, 266
Meckel, works cited, 273, 323
Median nerve in Raven, 125
Megalfema asiatica, 239, 240
Membrana nictitans in Raven, 57
MenobranchuB, 277

tibialis anticus in, 214

Menopoma, 277

mylo-hyoideus muscle in, 26

Merrem, work cited, 324
Method of exposing muscles of
upper extremity in Raven
for dissection, 66, 67
Method of removal of podotheca
of toes and tarso-metatarsus
to examine tendons, 155

336

INDEX.

Method of studying tensor patagii

muscles, 109
Menrsinge, work cited, 324
Mid-cricoidal segment of upper

larynx in Eaven, 45
Minot, work cited, 324
Mivart, on muscles and tendons in

legs of birds, 225, 226
works cited, 208, 214, 220,

261, 324
Monkeys, caudal muscles in, 261
flexor longus pollicis muscle

in, 141
Mourning Dove, bicipital slip to

patagium in, 109
Miiller, works cited, 324
Muscle, abductor mininddigiti, 140,

147, 151, 152
accessory femoro-caudal, 185,

186, 188, 231
accessory semitendinosus, 155,

167, 179, 188, 202, 224
adductor longus, 68. 155, 167,

194, 195, 196, 197, 224, 231
adductor niagnus, 68, 155, 167,

195, 197, 201, 224, 231

■ ambiens, 171, 172, 173, 176,

188, 231
anconeus, 83, 124, 126, 131,

133, 134, 143, 145

appendico-costales, 263, 293

biceps, 68, 77, 83, 89, 108, 109,

114, 116, 117, 121, 122, 126,

139, 140
biceps flexor cruris, 155, 158,

174, 175, 178, 184, 200, 207,

213, 216, 221, 224, 227, 229,

231, 260
biventer cervicis, 19, 260, 263,

270, 271, 273, 274
biventer maxillaj, 7, 15, 17, 18,

19, 23, 28
brachialis anticus, 77, 83, 89,

116, 121, 126, 132, 139, 145

broncliialis anticus, 43, 49, 50

broncliialis posticus, 43, 48, 50

broncho-trachealis anticus, 43,

47, 48, 50

broncho-trachealis brevis, 43,

48, 50

broncho-trachealis posticus, 43,

47, 48, 50

cerato-glossal, 24, 30, 46

cerato-hyoideus, 24, 27, 28, 31

circumconcha, 3,4, 71,5,28, 62

Muscle, cleido-trachealis, 3, 5, 6, 9,

14, 31, 46

complexus, 19, 262, 263, 266,

268, 269, 274

constrictor glottidis, 31, 43, 45

■ coraco- brachialis, 31, 80, 83,

95, 96, 97, 102, 107
coraco-humeralis, 68, 80, 83,

86
■ crura3us, 155, 170, 171, 178,

216, 221
■ deltoid, 31, 83, 87, 90, 97,

103, 108, 116, 117, 118, 119,

120
depressor coccygis, 196, 263,

300, 313, 314
depressor cauda), 68, 263, 293,

312

■ depressor-glossus, 24, 29, 30, 46

depressor palpebra) inferioris,

53, 56,61

dermo-cleido dorsalis, 3, 8 31

dermo-dorsalis, 3, 6, 8, 12, 14

dermo-frontalis, 3, 4

dermo-humeralis, 3, 14

dermo-iliacus, 3, 12. 260

dermo-pectoralis, 3, 14

dermo-spinalis, 3, 11, 260

dermo-temporalis, 3, 5, 7, 10,

15, 19, 28

dermo-tensor patagii, 3, 6, 7, 8,

10, 89, 112, 113!; 114

dermo-ulnaris, 3, 12, 68,78, 93

diaphragm, 263, 306, 307

digastric, 15, 17, 21, 28, 31, 46

entotympanicus, 15, 17, 19

expansor secundariorum, 78

79
extensor brevis annularis, 237

256
extensor brevis digitorum, 220.

231, 237
extensor digitorum communis

83, 121, 124, 125, 127, 131

148
extensor femoris, 155, 158, 166

167, 169, 173, 175, 178, 179

185, 187, 188, 216, 221
extensor hallucis brevis,

218, 220, 223
■ extensor indicis hmgus,

124, 131, 135, 143, 145

extensor longus digitorum, 1 56

178, 179, 187, 215, 218, 224,
231

156

121

INDEX.

337

Muscle, extensor metacarpi radialis

brevior, 134
■ extensor metacarpi radialis

longior, 83, 97, 114, 115,

116, 118, 121, 123, 124,

125, 127, 131, 133, 140, 147,

148
— — extensor ossis metacarpi jiol-

licis, 124, 126, 131, 132, 143,

145, 152
• — ■ — extensor proprins pollicis, 140,

147, 152
• femoro-caudal, 68, 155, 167,

183, 186, 188, 196, 224, 231,

293, 312, 317

flexor brevis hallucis, 237, 256

flexor brevis pollicis, 131, 140,

147, 148, 152
flexor capitis inferior, 17, 262,

266, 267
flexor carpi nlnaris, 83, 121,

124, 131, 140, 141, 142, 144,

152
flexor carpi ulnaris brevior,

124, 126, 140, 144, 145, 146,

152
flexor digitorum profundus,

121, 124, 126, 132, 139, 140,

141,142,144,145,146
flexor digitornm sublimis,

137
flexor longns hallucis, 156, 167,

178, 216, 221, 224, 231, 232,

234, 235, 237, 239, 240, 241,

251, 254
flexor metacarpi brevis, 121,

131, 147, 151
flexor metacarpi radialis, 83,

121,124,126,128,129,131
flexor minimi digiti, 121, 131,

147, 148, 152
• flexor minimi digiti brevis,

140, 147, 149, 152 _
flexor perforans digitorum

pedis, 227
-- — flexor perforans digitorum pro-
fundus, 156, 167, 137, 206,

234, 236, 237, 238, 239, 240,

246, 253
flexor perforatus annularis

primus pedis, 156, 221, 224,

241,243, 251, 252
flexor perforatus indicis primus

pedis, 156, 178, 179,206,221,

224, 231, 245, 253

Muscle, flexor perforatus indicis se-

cundus pedis, 156, 206, 216,

221, 229, 244, 253
flexor perforatus medius

primus pedis, 156, 209, 224,

242, 252
flexor perforatus medius secun-

dus pedis, 156, 187, 206, 216,

221, 242, 243, 246, 252, 253
flexor perforatus primus pedis,

243
gastrocnemius, 155, 175, 178,

179, 187, 197, 200, 207, 210,

213, 230
• gemellus, 68, 155, 178, 191, 194,

236, 237

genio-liyoideus, 24, 27, 28, 31

■ gluteus medius, 68, 155, 158,

160, 164, 178, 191, 216, 221,

260
■ gluteus minimus, 155, 163. 178,

185, 187, 216,221,260'
■ gluteus primus, 68, 155, 157,

158, 163, 171, 174, 207
infrjicoccygis, 196, 263, 300,

315, 317
■ interarticulares, 263, 280, 292,

294

intercostal es, 263, 296, 300

- — ■ interosseous dorsalis, 121, 131,

147, 153, 154
interosseous paimaris, 121, 131,

147, 153, 154
interspinales, 263, 269, 280,

291, 292
intertransversales, 263, 269,

274, 280, 294
lateralis caudte, 196, 263, 293,

314
• lateralis coccvgis, 196, 263, 300,

315, 317

latissimus dorsi, 80, 81, 82, 83,

97, 103, 118,260

levator caudae, 260, 263, 31 1

levator coccygis, 260, 263, 293,

307, 311
levator palpebral superioris, 53

55, 56, 61
levator scapula), 31, 80, 98, 99

104, 107, 260
levatores costarum, 263, 293,

297, 298, 300
longissimus dorsi, 260, 263,

269, 272, 276, 277, 278, 279,

281, 282, 283, 284, 293, 307

Z

338

INDEX.

Muscle, longus colli anterior, 263,
266, 274, 285, 286, 287

longus colli posticus, 260,

263, 269, 271, 272, 273, 274,
281, 282, 283, 292, 294

masseter, 7, 11, 15, 16, 17, 18,

19, 23, 28
mylo-hyoideus, 21, 22, 24, 27,

31, 32, 46
obliquo-traiisversales, 263, 269,

280, 294

obliquus colli, 263, 269, 273,

274, 281
■ obliquus externus abdominis,

263, 293, 301
obliquus interior, 53, 55, 56,

58,60
obliquus internus abdoniims,

263, 300, 303
obliquus superior, 53, 55, 56,

58, 59, 60
obturator externus, 68, 155,

185, 186, 188, 195, 197, 224,

231
-obturator internus, 155, 190,

191, 192, 194, 196, 236,

237
orbicularis palpebrarum, 53,

56, 61
pectoralis major, 14, 31, 67, 68,

69, 71, 77, 83, 86, 87, 89,

100, 103,108, 115, 302
pectoralis minor, 100, 101

pectoralis secundus, 67, 68, 71,

72, 77, 83, 87, 89, 103
■ pectoralis tertius, 67, 68, 71, 74,

78, 83, 89
peroneus longus, 156, 187,207,

208,209, 210,211, 212, 213,

214

plantaris, 204, 205

platysma mvoides, 3, 6, 7, 21,

23

popliteus, 237, 249, 250, 255

pronator brevis, 83, 121, 124,

129, 132, 140
^ pronator longus, 83, 121, 124,

132, 140
pterygoideus externus, 7, 15,

17, 20
pterygoideus internus, 7, 15,

17, 20, 23, 28

pvramidalis nictitantis, 53, 55,

"57, 60,61
— — quadratus lumborum, 299

Muscle, quadratus nictitantis, 53,55,

57, 60, 61
■ rectus abdominis, 263, 300,

303, 304
rectus capit'8 anticus minor,

19, 262, 265, 266, 267, 268,

274
— — rectus capitis lateralis, 17, 263,

269, 274, 289, 290
rectus capitis posticus major,

19, 262, 268,269,271, 274
rectus externus, 53, 55, 56, 59,

60
rectus inferior, 53, 55, 56, 59,

60

rectus internus, 53, 55, 56, 60

rectus superior, 53, 55, 56, 59,

60
rhomboideus, 31, 68, 80, 84, 97,

103
sacro-lumbalis, 263, 269, 276,

287, 278, 281, 282
sartorius, 68, 155, 156, 157, 158,

172, 207, 213
scalenus medius, 263, 293,

297
scapulo-humeralis, 31, 86, 80,

83, 87, 103
semimembranosus, 68, 155, 177,

180, 181, 207, 213, 216, 221,
224, 231

semitendinosus, 155, 177, 179,

181, 188, 195, 207, 213, 216,
221, 224, 231, 260

serratus magnus anticus, 68, 80,

92, 93, 99, 104 '
serratus parvus anticus, 31, 68,

80, 104, 107
soleus, 156, 187, 204, 205, 206,

208, 236, 237, 247, 255

sterno-hyoideus, 24, 29, 31, 46

sterno- tracheal is, 43, 47, 49, 50,

51
• subclavius, 31, 71, 75, 80, 87,

94, 96, 107
subscapularis, 31, 80, 83, 87,88,

96, 101, 102, 107
supinator brevis, 83, 124, 126,

127, 128, 143, 145
supraspinal us, 68, 80, 83, 88,

91, 92, 103, 120
stylo-hyoideus, 7, 23, 26, 28,

31
■ temporal, 5, 7, 11, 15, 16, 17,

18, 19, 28

INDEX.

339

Muscle, tensor patngii brevis, 31, 77,

83,89,97, 105, 106,112, 114,

123, 125, 131, 140
tensor patagii longus, 6, 8, 31,

77,89,97, 105,106, 108,109,

112, 114, 115, 121, 123, 124,

131

r tensor tynipani, 53, 56, 62

teres et infraspinatus, 68, 80,

83, 90, 91, 92, 97, 98, 103,

120
teres minor, 31, 80, 83, 96, 97,

98, 102, 107
thoraco-scapularis, 68, 80, 99,

100, 101, 102, 104, 107
tliyreo-arvtenoidens, 31, 43, 45,

46
tibialis anticus, 156, 167, 179,

187, 201, 207, 209, 211, 212,

213, 214, 216, 217, 219,

221
• tibialis posticus, 156, 178, 216,

221, 224, 227, 228, 229,

231

tracheo-lateralis, 43, 47, 50

frachelo-mastoideus, 17, 263,

269, 274, 289, 290
transversalis abdominis, 263,

300, 305, 307
transversus perinei, 68, 263,

293, 312, 314, 317
• trapezius, 31, 68, 80, 82, 97,

103, 108
■ triangularis sterni, 263, 295,

300, 306
triceps, 68, 77, 83, 89, 91, 103,

114, 116, 119, 120, 121
vastus externus, 155, 167, 170,

171, 178, 185, 216, 221
vastus internus, 155, 167, 172,

173, 175, 236, 237
Muscles, characters of, in birds, 1

■ method of dissecting, 1 5

of forearm and hand of Raven,

122

• of hand in Raven, 147

of the brachium in Raven,

116

of the head, 15

of the lower extremity, 155

Musculi accessorii ad sacro-lum-

baleni of Owen, 278
Myological formula) of Garrod for

muscles of the thigh in birds,

188

N

Natatohes, coraco-bracliialismuscle

in, 9(3

pectoralis ninjor muscle in, 71

Neander, work cited, 324
Nitzsch, work cited, 324
Nitzsch-Giebel, works cited, 324
Nycticebus, flexor longus pollicis

muscle in, 141
soleus muscle in, 208

O

Obturator foramen, 193

Obturator space, 236

CEsophagus, action of digastric

muscle on, in deglutition, 22
Olecranon process, 139, 141, 142
Oiieration of muscles of air pas-
sages in Raven, 51
Optic nerve, 57, 60
Order in which muscles of forearm

should be examined, 146
Ornithorhynchus, gemellus muscle

absent in, 194

■ soleus muscle in, 208

Os furcula, 10

Os humero-scapulare, 83, 103, 118

Os magnum, 146, 149

Os prominens in Hawks and Owls,

108, 109
Os radiale, 115, 146
Osseous plate at back of eye in

Raven, 60
Ostrich, absence of lower larynx in

49
■ flexor metacarpi radialis muscle

in, 129

flexor metacarpi ulnaris in, 144

pectoralis major muscle in, 71

tensor patagii muscles in, 107

Owen, Sir Richard, on caudal muscles

in the Apteryx, 316
on muscles and their tendons

in legs of birds, 225, 226
sterno-maxillaris muscle of, 9,

10
works cited, 210, 211, 214, 215,

219, 220, 259, 325
Owl, organ of hearing in, 44, 45

tensor tympani in, 62

Owls, bony bridge on tarso- metatar-
sus in, 218
semitendinosus muscle absent

in, 181

340

INDEX.

P

Parapatagium, 7
Parker (T. J.), work cited, 325
Parker (W. K.), works cited, 32.5
Parrot, lower larynx in, 44, 45, 51

the tonf,nie in, 52

Pars metapatagialis of the M. ser-

ratus superhcialis .■<. thoraci-

scapularis of Fiirbringer dis-
cussed, 12
Pars propatagialis musculi cucnl-

laris, 8, 10
Passeres, tibialis posticus in, 229

vinculum absent in, 235

Patella, 158, 159, 171, 17(3

Pelican, upper larynx in, 47

Pelvic limbs in Cypseli and Trochili,

190
Pelvis, 14

post-pubic elements of, 14

Penguin, extensor metacarpi radialis

longior muscle in, 124
flexor metacarpi radialis muscle

absent in, 129

flexor metacarpi ulnaris in, 144

pectoialis major muscle in, 71

Penguins lack accessory semitendi-

nosus, 181
pectoralis secundus muscle in,

73, 74

pectoralis tertius muscle in, 76

Pericardium, relations to diaphragm

in Raven, 306
Peritoneum, 305
Perrin, work cited, 325
Petrels, pectoralis secundus muscle

in, 74
Pfeiffer, work cited, 325
Phalacrocorax carbo, longus colli

posticus in, 275
Phrynosoma coronatum, 261
Pig-, soleus muscle absent in, 208

tibialis anticus in, 214

Pinon Jay, obturator internus muscle

in, 192
Plantar tendons in different birds,

238, 239, 240
Plotus anhinga, longus colli posticus

in, 273, 275
Plotus, longus colli in, 287, 288
Plovers, bicipital slip to laatagiuni

in, 109
expansorsecundariorum muscle

in, 79

Podiceps minor, semimembranosus
muscle in, 181

PoUex in Raven, extensors and flexor
devoted to, 148

Porpoise, coccyx of, 262

Posterior patagial fold, relations of,
67

Prechtl, work cited, 325

Pronator group of muscles of fore-
arm, synonymy of, 129, 130

Propatagiuni, 7

Purple Martin, patagial muscles in,
113, 114

Pygostyle, 184

Q

Quadrate bone, masseter attached
to, 17

orbital process of, 20

relations of certain muscles to,

19

Quadrato-jugal bone, origin of ten-
sor tympani, 62

Quennerstedt, work cited, 325

R

Rails, expansor secundariorum

muscle in, 79
Raptores, pectoralis secundus

muscle in, 74
Raptorial birds, biventer cervicis

in, 272
Rasores, coraco-brachialis muscle

in, 96

pectoralis major muscle in, 71

upper larynx in, 47

Raven, muscles of the trunk of,

262

patagial muscles in, 114

preparation for dissection of, 1

Ray, work cited, 325
Recapitulation of eye-muscles in

Raven, 61
Rectus feiiioris, of Owen (Apteryx),

158
Reid, work cited, 325
Reptiles, digastric muscle in, 22

longissimus dorsi in, 279

ventral muscles in, 262

Retzius, work cited, 325

Rhea, tensor patagii muscles in, 107

RhyncliBea, air passages in, 52

INDEX.

341

Rima glottidis in Raven, 45
Rings of trachea in Raven, 43
Rolleston, work cited, 325
Roof of mouth, formed largely

by iDterygoideus internus

muscle, 20
Ruedinger, work cited, 326

S
Sabatter, VFork cited, 326
Salter, Dr., named the cerato-glossus

muscle, 33
Sanders (A.), work cited, 261
Savart, on sterno-trachealis muscle,

49
Schneider (A.), work cited, 326
Schneider (J. G.), work cited, 326
Schopss, work cited, 326
Sciatic artery, relation of accessory
femoro-caudal muscle and
obturator externus, 185, 188
Sciatic nerve, relation of accessory

femoro-caudal muscle, 185
Sclerotal plates, 58
Seal, soleus muscle absent in, 208
Secondary remiges in Progne subis,

114
Selenta, on the dermal muscles, 3

. Avorks cited, 326

Sesamoid of foot of Raven, 178
Shoulder-girdle, muscles attached to,

31
Shufeldt, works cited, 326
Snakes, muscles of the dorsum, 259
Stannius, work cited, 326
Steno, work cited, 326
Strigidae, eye closed by upper lids

in, 57
Structure of windpipe in Raven, 43
Struthionidae, coraco-brachialis

muscle absent in, 96
Struthious birds, pectoralis secundus
muscle in, 74

trunk muscles in, 259

Sula fusca, longus colli posticus in,

275
Sundevall, works cited, 235, 327
Superior larynx in various birds, 47
Sutton, references to expansor secun-
dariorum muscle ; compares
with coraco-brachialis longus
of mammals, 79, 80

work cited, 327

Swallows, pectoralis major muscle
in, 71

Swans, expansor secundariorum

muscle in, 79
Swifts, not especially related to the

Humming-birds, 189
Synonymy of adductor muscles of
thigh in birds, 194, 195

anconeus muscle, 134

biceps flexor cruris, 174

biceps muscle, 116

biventer cervicis, 270

brachiaHs anticus muscle, 122

caudal muscles in birds, 308,

309, 310, 311

complexus, 263

coraco-humeralis muscle, 86, 95

deltoid muscle, 117, 118

expansor secundariorum

muscle, 79

extensor digitorum communis

muscle, 125

extensor femoris, 169, 170

extensor hallucis brevis, 218

extensor indicis longus muscle,

135

extensor longus digitorum, 215

extensor metacarpi radialis

longior muscle, 124

extensor ossis metacarpi pollicis,

132

extensor proprius pollicis, 147

femoro-caudal, 183, 184

flexor brevis pollicis, 148

flexor capitis inferior, 267

flexor carpi ulnaris, 141

flexor carpi ulnaris brevior,

notes on, 144

flexor digitorum profundus,

139

flexor digitorum sublimis

muscle, 137, 138, 139

flexor longus hallucis, 232

flexor metacarpi brevis, 151

flexor metacarpi radialis, 128

flexor minimi digiti, notes on,

148
flexor perforans digitorum pro-
fundus, 247

flexor perforans et perforatus

digiti II. of Gadow (flexor
perforatus indicis secundus
pedis), 229, 230

flexor perforans et perforatus

digiti III. of Gadow (flexor
perforatus medius secundus
pedis), 243, 244

342

INDEX.

Synonymy of flexor perforatus diiiiti
11. of Gadow (flexor perfor-
atus indicis primus pedis),
245, 24'i

flexor perforatus digiti III of

Gadow (flexor perforatus
medius primus pedis), 242,
243

flexor perforatus digiti IV. of

Gadow (flexor perforatus an-
nularis primus pedis), 241

gastrocnemius, 200

gemellus muscle, 194

gluteal muscles, 159, 160, 161,

162, 163, 164, 165, 166

intercostales, 296

interosseous dorsalis, 153

interosseous palmaris, 153, 154

intervertebral muscles, 291, 292

latissimusdorsi, 81

levator scapula3 muscle, 98, 99

levatores costarum, 298

lingual muscles, 34, 35, 36, 37,

38, 39, 40, 41, 42

longissimus dorsi, 278

longus colli anterior, 288, 289

longus colli posticus, 276

muscles of dorsum in birds,

277, 278, 279

muscles of eye and ear, 63, 64,

65

mvlo-hyoideus, 24, 25

obliquiis colli, 281, 282

obliquus externus abdominis,

301

obliquus internus abdominis,

303

obturator externus muscle, 186

obturator internus, 190, 191

pectoral muscles, 69

pectoralis secundus muscle, 72

pectoralis tertius muscle, 74

peroneus longus, 208, 209

peroneus profundus of Gadow

(tibialis posticus), 228, 229

popliteus, 249, 259

pronator brevis muscle, 129

rectus abdominis, 303

rectus capitis anticus minor,

266

rectus capitis posticus major,

268

rbomboideus muscle, 84

• sartorius muscle, 156

scalenus medius, 297

Synonymy of scapulo-humeralis
muscle, 86

semimembranosus muscle, 180

semitendinosus, 177

serratus magnus anticus, 92

soleus (plantaris ?) muscle, 205

subclavius muscle, 94

subscapularis muscle, 101

-supinator brevis muscle, 127

• suprasjjinatus muscle, 88

teres et infraspinatus, 90

tensor patagii muscles, 105

thoraco-scapularis muscle, 92,

99

tibialis anticus muscle, 211, 212

traclielo-mastoideus, 289, 290

transversalis abdominis, 305

transversus perinei, 312

trapezius muscle, 82

■ triangularis sterni, 295

triceps muscle, 119

T

Tarsal cartilage of eye in Raven, 54

of leg in Raven, 178

Teleosts, muscles of tbe dorsum, 259
Tendinous loop connected with biceps

of the thigh in birds, 175
Tendinous raphe of accessory semi-
tendinosus, 179
Tendons, characters of, in birds, 1

■ tendency to ossify in birds, 1

Tensor fasciae, of Garrod, 158
Tensor patagii muscles in taxonomy,

106
Tensor vaginae, of Cuvier, 158
Tensor vagina;, of Meckel, 158
Tensor vagina;, of Owen (Apteryx),

158
Thamnophilus, sterno-trachealis

muscle in, 51
Thigh-muscles of Trochili and

Cypseli, 189
Thuet, work cited, 327
Thyroid plate, 45
Tibial cartilage, 209, 210, 230, 232,

236, 237, 245, 248

ossification in, 208

Tiedemann, work cited, 327
Tongue, as affected by action of

stylo-hyoideus muscle, in

Raven, 27
as affected by contraction of

mylo-hyoideus muscle, in

Raven, 26

INDEX.

343

Tongue, as affected by the cerato-

hyoideiis muscle, in Raven,

29
as affected by tlie genio-

hyoideus muscle, in Raven, 27
as affected by the sterno-

hyoideus muscle, in Raven,

30

muscles of, in Raven, 23

Touraco {Corythaix eri/throlopha),

ambiens muscle in, 171, 173
Trachea, 10, 22, 43, 48, 49
Trunk, musculature of, 258
Tubercle on tarso-metatarsus for

tibialis anticus, 217
Tubinares, patagial muscles in, 114

pectoralis tertius muscle in, 76

Two-toed Anteater, teres minor and

infraspinatus muscles in, 98
Tympanic membrane in Raven, 50

U

Ulnar tuberosity of humerus,

muscles attached to, 91, 96,

97, 102
Ulnare of carpus, 142
Ulrich, work cited, 327
Under side of eye in Raven, its

muscles, 60
Upper extremity, muscles of, 66
Upper larynx, 6, 10
Upper side of shoulder-joint, muscles

attached to, in Raven, 85
Upupa epops, 235
Urodeles, tibialis anticus in, 214

Veins, gastric, in Apteryx, 307

Vertebra dentata, 272

Vertebral ribs, 13

Vertebrata, tibialis anticus muscle

in, 214
Vetter, work cited, 327
Viallane, temporo-alaris muscle of,

6

work cited, 327

Vicq d'Azyr, work cited, 327

Vinculum, 234, 235, 238, 239
Voice, where formed in birds, 47
Volitores, muscles of air passages in,

51
Vulture, absence of lower larynx in,

49
Vultures, diaphragm in, 307

W

Wagner, work cited, 327
Waller, work cited, 327
Watson, works cited, 327
Weitzel, work cited, 327
Weldon, work cited, 327
Wiedemann, work cited, 328
Wing, closing and spreading of, in

birds, 148
AVoodpecker, Owen's cerato-tra-

chealesin, 33
tongue a.nd salivary glands in,

44
Woodpeckers, subvertebral caudal

muscles in, 262
Work to be done on thigh-muscles

in birds, 190

X, SYMBOL for semitendinosus, 188
Xiphoidal border of sternum in
Cypseli and Trochili, 190

Y, SYMBOL for accessory semitendi-
nosus, 188
Yarrell, works cited, 328
Young, work cited, 328

Zoological Society of London, Pro-
ceedings of, Forbes's con-
tributions to, 1 14

Zootomy (Parker's), tensor patagii
accessorius of Common
Pigeon in, 110

Zygoma, 11, 17, 18, 19, 108

THE END.

Richard Clay and Sons, Limited,
london and bungay.

Iff