Jollie 1957 Chick Skull

T H E HEAD SKELETON O F T H E CHICKEN AND
REMARKS ON T H E ANATOMY O F THIS

REGION I N OTHER BIRDS
MALCOLM T. JOLLIE
Department of Biological Sciences, University of Pittsburgh,
Pittsburgh 19, Pennsylvania
TWENTY-SIX FIQUBEB
INTRODUCTION
A justification for another account of the bird head skele-

ton lies in the fact that earlier ones are either so complex
(Parker, 1869)’ so brief and general that it is difficult to
identify some of the parts (Newton and Gadow, 1893-1896;
Stresemann, ’27-’34; GrassQ, ’50)’ so confused in their ter-
minology that the identity of some bones is in question (Mag-
nus, 1871)’ or just incomplete (Schinz and Zangerl, ’37).
Usually it is a combination of these factors that leaves one
in doubt (Marinelli, ’36; Erdmann, ’40).
An example of the existing confusion is presented by the
figure of Heilmann ( ’26, fig. 3). He shows the opisthotic as
forming the posterior margin of the tympanic cavity in the
goose (Amer) while Stresemann (’27-’34, fig. 53)’ using the
same drawings, has united Heilmann’s exoccipital and opis-
thotic and labelled the combined bone the exoccipital. Ihle
(’47, fig. 227) has used this same figure but introduced some
new labels. Not only is there confusion of parts but the exact
extent of various bones and their interrelationships with other
bones cannot be determined from the available descriptions ;
this is particularly true of the bones of the auditory capsule,
the basis cranii and the posterior wall of the cranium.
New methods of study have appeared since the early foun-
dation works on vertebrate skulls; the most important of
389
JOUENAL OF MOEPHOLOGY, VOL. 100. NO. 3
MAY 1957
390 MALCOLM T. JOLLIE
these is the use of whole, cleared specimens the bones of which

have been stained with alizarin. This technique has been
reviewed and described by various authors (see Petri, '35).
This type of study has been used in studying the development
of the bird skull by Schinz and Zangerl ( '37 -chicken, pigeon
and grebe). Huggins, Huggins, Hellwig and Deutschlander
( '42 -House Wren) and Maillard ( '48 - Skua) .
This paper is a part of a general review of the dermal and
chondral ossifications in the heads of the different classes
of vertebrates the basic purpose of which is to "homologize"
the bones of the different classes. The homologizing of bones
is fraught with many difficulties and strong differences of opin-
ion are encountered, but advances in our knowledge of the
fossil record and a clearer understanding of the structure
of some of the more primitive living types have made it pos-
sible to more accurately determine the evolutionary history
of each unit.
An important key to the phylogeny of a bone lies in its
ontogeny. Here again new information and methods produce
more understandable and useable results. Also in this regard,
Smith ('47) has pointed out that some of the usual concepts
of bone origins are not necessarily accurate. A consideration
of the development of the bird skull supports his views.
If in spite of our broadening knowledge of head skeletons
we cannot agree on homologies, at least we can arrive at a
more consistent terminology.
I want to thank Dr. Charles Peterson and the members of

the Poultry Department of the University of Idaho f o r sup-
plying the specimens used in this study and the National
Science Foundation for its financial support.
This study is based primarily on heads of white leghorn
chickens stained with alizarin and cleared in glycerin (see
Evans, '48). I n age the specimens range from those showing
the first evidences of ossification (at 10 days of incubation)
to those in which the bones of the cranium are thoroughly
HEAD SKELETON O F CHICKEN 391
fused (100 days or more after hatching). Exact ages cannot

be given for incubation specimens since the rate of develop-
ment and the starting point of development cannot be ascer-
tained (Erdmann's, '40, time designations cannot be con-
sidered as accurate-his specimens of 9 days look much
like mine at 10-11 days and probably correspond in stage
of development).
The stained and cleared type of preparation has many
advantages over study of serial sections. F o r example, the
slightest ossification stands out sharply and the shape and
relationships of each bone can easily be determined ; in addi-
tion, large numbers of specimens can be compared. The
difficulties of studying serial sections, combined with the fact
that the really critical changes occur rapidly in the period
from the 12th to the 17th days of incubation, accounts for
our present rather hazy notions. As a check, and for histo-
logical detail, a single skull (12th day of incubation) was
sectioned at l o p and stained with Mallory's triple stain.
There might be some doubt as to the accuracy of the alizarin
staining method but comparisons with similar stages studied
by the serial method dispells this. From all evidence it is
completely reliable and much less subject to error than at-
tempting reconstructions. The alizarin method detects bone
somewhat later than the x-section method, where the massing
of mesenchyme (blastemata) can be observed well before any
actual ossification occurs.
The development of the chick head skeleton was compared
particularly with stages of that of the Mallard Duck ( A m s
platyrhynchos [Linnaeus] ) , Cooper Hawk (Accipiter cooperii
[ Bonaparte] ), Golden Eagle (Aquilo chrys&tos [Linnaeus] ),
Magpie (Pica pica [Linnaeus] ), American Robin ( T u r d u s
migrntorius [Linnaeus] ) , English Sparrow (Passer domesti-
cus [Linnaeus] ), House Finch (Carpodacus mesicanus [P. L.
S . Miiller]), and Oregon Junco ( J u m o oregolzus [J.K. Town-
send]) and with the accounts in the literature (particularly
Parker, 1869 ; Magnus, 1871; Tonkoff, '00 ; deBeer, '37 ; Mari-
nelli, '36; Schinz and Zangerl, '37; and Erdmann, '40). The
foramina of the chicken skull are described and figured by

Kesteven ( '25).
The head skeleton of the bird in most features is typically
reptilian but differs strikingly in several ways. The general
proportions are altered as a result of an enlarged brain and
eye. The reptile has room f o r large fossae outside the brain
capsule ; this is not the case in the bird. Rounding of the bird
cranium is sometimes accentuated by the thickened spongy
(diploe) character of the bone. The interior spongyness of
pREMnXlLLA, x
EXOC OCCIPITAL
- QUAMATE
Fig. 1 Lateral view of 2-3 day old chick head skeleton.
the bone has allowed penetration by air sacs, particularly in

the tympanic region (see Bremer, '40).
The general resemblance of the pattern of bones in the
bird head skeleton to that of the reptile suggests that a direct
comparison can be made - such a comparison demands few
changes in prevalent terminology and these changes have been
suggested, or inferred in other accounts. The order of de-
scription will be from the posterior to the anterior end, without
separation of the membral and chondral bones. The following
comments apply to the chicken unless otherwise indicated.
HEAD SKELETON OF CHICKEN 393
SKULL
Basioccipital (figs. 3-10, 13)

The basioccipital is a ventral midline, chondromembral bone
(see Smith, ’47, on this designation) which supplies the central
part of the occipital condyle. It appears early in the 13th
day as an ossification around the ventral half of the notochord,
just anterior to the occipital condyle. I t gradually increases
in dimensions to enclose a part of the notochord, and is not
PREFRONTAL
NASAL I
PRAWCIPITAL
Fig. 2 Dorsal view of 2-3 day old chick skull.
completely developed until its fusion with the surrounding

bones (about 90 days after hatching). The basioccipital is
fairly thick, particularly anteriorly where it abuts along an
oblique line (as viewed in sagittal section) with the basi-
sphenoid. The ventral aspect is hollowed out just anterior
to the condylar process. Laterally it abuts against the ex-
occipital. The ventral tip of the lagena lies between its
anterior end and the exoccipital.
Esoccipital (f;gs. 1, 3-10, 12, 13, 17)

An exoccipital lies to either side of the foramen magnum.
These bones arise as perichondral ossifications (chondro-
membral) early in the 12th day of incubation and gradually

expand outward from the first area of ossification. The medial
margin is indented for the several divisions of the hypoglossal
nerve (XII). Laterally, extensions from the exoccipital form
pUADRATw- ,QUADRATE
\ -
Fig, 3 Veiitral view of 2-3 day old chick skull.
the posteroventral and posterior margins of the tympanic

cavity- this is also the case in the other kinds of birds
examined. On its internal aspect the exoccipital bears the
opisthotic, with which it becomes indistinguishably fused soon
after the appearance of that bone (figs. 6, 9, 10).
Each exoccipital contributes a small part to the occipital

condyle. It abuts anteriorly against the basisphenoid, dorso-
laterally with the squamosal, posteromedially with the supra-
occipital and ventromedially with the basioccipital. It is pene-
trated by foramina for the hypoglossal, spinal accessory,
Fig. 5 Median view of 2-3 day old chick skull, cut in half sagittally, along
with median view of mandible.
PROOTK:
IC
Fig. 6 Dorsal view of cranial cavity and rostrum of chick skull at the 16th
day of incubation.
vagus and glossopharyngeal nerves as well as the blood ves-

sels. I t is underlain dorsolaterally by an air space from the
middle ear cavity, extensions of which penetrate the parietal
and supraoccipital bones.
Supraoccipital (figs. 1-10, 12)

The supraoccipital usually arises from two perichondral
centers but W. K. Parker (1866: 124; 1873: 105) found only
one in Turdus. Gadow and Selenka (1891: 18) list the dove
and swallow as having only a single, medial center; T. J.
r 8
SUPRACCCIPITAl
9 10
Fig. 7 Occipital bones of the chick skull at the 14th day of incubation. The
bones are laid flat around the foramen magnum.
Fig. 8 As in figure 7, but at the 15th day of incubation
Fig. 9 As in figure 7, but late in the 15th day of incubation.
Fig. 10 As in figure 7, but at the 16th day of incubation.
Parker (1891 : 67) found only one in A p t e r y x , Sushkin (1899 :

80) reports only one for Falco tinnurzcuulzrs Linnaeus and
Maillard ( '48 : 76) describes only one for the Skua (Catharacta
skua Briinnich). The Magpie, English Sparrow, and House
Finch appear to have only one center but the bilateral centers
probably merge before ossification begins.
The two ossification centers of the chicken appear during
the 14th day of incubation (figs. 7-10) and soon fuse at the
midline (15th day of incubation) to form a plate which lies
above the foramen magnum. This plate has a notch in its
SEMICIA. WNAL
SEMICIA CANAL
\
BAsmcIPITAL
Fig. 11 Occipital bones of the English Sparrow about one week after hatching.
dorsal margin at the line of fusion. I n the chick the supra-

occipital almost spans the arch between the exoccipitals, but
in the hawk (Accipiter, Aquiln), it is a narrow vertical plate
with a midline notch. A narrow supraoccipital, such as that
of the English Sparrow, leaves the epiotic exposed lateral
to its margin (fig. 11).
The posterior wall of the cranium may have two large
fontanelles, one on either side of the supraoccipital; the
pigeon (Columbn Zivia Gmclin) is peculiar in that it has a
small, single, midline fontanelle. Where bilateral openings
are present, as in the Mallard Duck, a narrow band of the
supraoccipital extends laterally below the fontanelle to fuse
with the epiotic. Such a lateral extension is indicated in
Falco tinm.mcu1u.s (Sushkin, 1899, Taf. V, fig. 95), a species

in which fontanelles do not occur in the adult. I n most birds
there is a small foramen, or two, to either side of the supra-
occipital through which a vein emerges from the cranium.
It is of interest that although the cranium of the chicken is
not particularly inflated or rounded as compared with that
of many birds (hawk, passerine) yet its supraoccipital is
broad while that of the others is very narrow. Expansion of
the cranial walls appears to be correlated in part with medial
compression of the supraoccipital, although a narrow cranium
goes along with a narrow supraoccipital in NzLmenium ameri-
c a v w Bechstein.
In the chicken a diverticulum from the middle ear cavity
penetrates the lateral margin of the supraoccipital.
# # #
Associated with the occipital ring are the bones enclosing

the inner ear. In the tetrapod (and crossopterygian?) these
may be three in number :prootic, opisthotic and epiotic. Based
on its relationships to the inner ear, the prootic appears to
be a new element which has expanded to replace the sphenotic
and part of the pterotic of some bony fishes while the opis-
thotic is roughly comparable to that of the fish and has invaded
a part of the region of the pterotic. The use of these terms
must be considered purely provisional. It would probably
be much more proper to call the prootic the petrosal and the
opisthotic the mastoid bone (Gadow and Selenka, 1891 :24-25).
A separate epiotic is not known to occur in reptiles or am-
phibians (Williston, '25: 22; Romer, '55, does not mention
it). A separate opisthotic occurs only in Cotylosauria, Che-
lonia (Williston, '25: 23) and Mammals.
Prootic (Petrosal -figs. 1, 5, 6, 12, 13)

The prootic arises from a single chondromembral center
which appears late in the 14th day of incubation. This bone
covers the inner aspect of the labyrinth and encloses it except
for the middle portion of the anterior vertical semicircular
HEAD SKELETOR O F CHICKEN 399
canal (enclosed in the epiotic), the posterior vertical semi-

circular canal along with its ampulla and much of the hori-
zontal semicircular canal (enclosed in the opisthotic), and
the tip of the lagena (between the basioccipital and exoccipi-
tal). The inner surface of the prootic is perforated by several
branches of the auditory nerve and by the endolymphatic
duct (fig. 5). The fenestra ovale, into which the columella
(stapes) fits, lies between the prootic and opisthotic. The outer
dorsal aspect has an articular facet for the inner head of the
ITAL
Fig. 12 Lateral view of the posterior end of the 2-3 day old chick skull with
the squamosal removed.
quadrate and is separated from the squamosal by an air sac

from the middle ear cavity. The form of the head of the
quadrate, or the distance between the squamosal and prootic
articulations, is related t o the size of this air space.
Opisthotic (Mastoid -figs. 1, 5, 6, 9, 10, 12, 13)
The chondromembral opisthotic appears early in the 15th

day of incubation as a center which is very irregular in out-
line (may be multiple) and which is associated primarily with
the outer surface of the ampulla of the posterior vertical
semicircular canal. It eventually covers the posterior aspect
of the membranous labyrinth, enclosing the posterior vertical
semicircular canal and its ampulla and a part of the horizontal

canal. An anterolateral splint rims the ventral half of the
fenestra ovale and the dorsal half of the fenestra rotundum
(margined ventrally by the exoccipital -fig. 13). The antero-
medial margin is notched by a foramen f o r the 9th and 10th
cranial nerves.
The question of whether-or-not the opisthotic remains ex-
posed in the posterior cranial wall cannot be decided with
complete satisfaction (see Erdmann’s, ’40 : 347-350, descrip-
ANT VERT SEMlClR CANA
POST VERT
SEMlClR CANA
HORIZONTAL CANA
LAGENAR OTOLITH
Fig. 13 An terolateral view of the ear area of the chick skull at the 16th day
of incuhal;ion ; the squamosal, parietal and supraoccipital not shotwn.
tion of this area). During the 16th day of incubation (figs.

7-10) the opisthotic of the chicken is exposed, but later it
appears to be covered by a sheet-like outgrowth from the
exoccipital (18th day of incubation) or at least its surface
ossification is directly continuous with that of the exoccipital
and appears to form as an extension from the margin of
that bone. This “covering” of the opisthotic takes place
much later in the English Sparrow, at about 10 days after
hatching. In the Mallard a part of the opisthotic is distinct
from the exoccipital; this suggests that there is a true external
ossification of this bone which is separate from the exoccipital
but later fuses with it (as in goose or Regadis-Gadoxv

and Selenka, 1891, Taf. 11, fig. 10). The fact that the opisthotic
fuses with the exoccipital from its beginning suggests that
this center is much repressed in the bird as it is in the reptile
and ( ?) mammal. The opisthotic probably never supplies a
point of articulation for the quadrate (as suggested by
Lowe, ’26).
Epiotic (figs. 5, 6, 9 )
The epiotic appears early in the 16th day of incubation
and soon encloses a part of the loop of the anterior vertical
semicircular canal. It may be fused to the ventrolateral inner
aspect of the supraoccipital (chick), or be exposed on the
posterior aspect of the skull as in Hirumdo (see Gadow and
Selenka, 1891: 26-27, Taf. 111, fig. lo), Turdus or Passer
(fig. 11). I n the Ostrich, Rhea, and Red-tailed Hawk (Buteo
jamaicelzsis [Gmelin]) it appears to give rise to a distinct
part of the posterior wall. Again, as in the case of the opis-
thotic, it is a question whether the supraoccipital spreads
out as a thin surface cover over this bone or whether the
surface ossification can truly be called epiotic. I am inclined
to accept the latter view.
# + +
The ossifications in the basis cranii present one of the
most difficult problems of the bird skull. A brief consideration
of this region in the reptile might help in understanding the
bird. According to Williston ( ’25: 26) the parasphenoid is
usually indistinguishably fused to the underside of the basi-
sphenoid in early amphibians and most reptiles, with the
exception of some gechos. It is much reduced in gechos, and
many lizards, or apparently lacking in crocodilians, turtles
and some lizards (Williston, ’25, figs. 30, 69; Goodrich, ’30,
figs. 415, 416, 420; McDoivell and Bogert, ’54, fig. 33; Rao
and Ramaswami, ’52, figs. 10, 10a). Parker (1880-1885 : 285,
pl. 66, fig. 3 ) describes investing “basitemporal” plates
lateral to the “basisphenoid,” which in section is a dermal
bone underlying the hypophyseal fossa, so that actually the

parasphenoid appears to be well represented in the crocodile.
The actual number of elements involved in the bird is not
clearly indicated. According to Parker (1890 : 6-7) there are
three ossification centers in the bird parasphenoid which fuse
to form the broad “basitemporal plate” and the median an-
terior rostrum. He believed that the parasphenoid replaces
the normal endoskeletal bony centers of the basisphenoid and
presphenoid and that the pituitary is enclosed ventrally by
this bone. Selenka and Gadow (1891: 20-22) and deBeer
(’37: 280-281) assumed that there are 5 centers: a rostral,
two basitemporals and two basisphenoids. They assumed
that the two latter pairs fused. Erdmann ( ’40: 357-364)
described 7 centers: one rostral, two basitemporal, two basi-
sphenoid and two tympanic wing centers of the basisphenoid.
Kesteven (’41: 332) had somewhat different views. I n 15
specimens he observed no dermal ossification centers ; rather
there was an ectochondral ossification of basisphenoid and
“presphenoid. ”
The possibility of a “presphenoid” being involved cannot

be disregarded (Kingsley, ’25 : 175 ; Stresemann, ’27-’34 : 49,
have described separate “presphenoid” centers). An un-
questionable “presphenoid” is known only for the mammal,
but even here it is only the fused midline portion of the orbito-
sphenoids according to Goodrich ( ’30: 395) who adds that
“There appears to be no separate ‘presphenoid’ bone in
any adult Mammal.” The origin of the mammalian orbito-
sphenoid is described by Kingsley (’25 : 188-189) - the “pre-
sphenoid” arises from a pair (or two) of centers while the
“orbitosphenoid” arise on either side from single centers ;
all of these centers fuse during the later development. I n the
human, the development of this part of the sphenoid is similar
(two pairs of centers -Gray, ’42 :173) ;in insectivores ( R o n ,
’47 : 370-376) the presphenoid part never arises from a single
median center.
The orbitosphenoid of the bird arises from dorsal and

ventral centers which can be homologized with the “orbito-”
and “pre-” sphenoids of the mammal. The origin of the
“pre-” sphenoid by the ventral fusion of the submammalian
orbitosphenoids seems the logical explanation of the sudden
appearance of this “new bone” in the mammal and thus
removes the possibility of its occurrence in the bird.
A histological examination of the developing basis cranii
of the bird (figs. 14-16-see illustrations of T. J. Parker,
1891, pl. 13, figs. 160-162 ; pl. 14,figs. 163-166) does not really
solve the problem of its formation since the chondral basi-
sphenoid of the mammal is not matched by the membral an-
lage of the bird basis cranii (refer to Smith, Copenhaver and
Johnson, ’53: 117, on bone development). T. J. Parker (1891)
has differentiated a membral parasphenoid from a membral
basisphenoid in a most arbitrary way. That ossification of the
trabecular cartilages does occur is quite apparent and the
name basisphenoid should be restricted to the perichondral
(membral) and later endochondral ossification of these car-
tilages. If this is done then in the chicken basis cranii the
basisphenoid can be seen only from above as it is limited
to a part of the lining of the selln turcica, its dorsal margin
and the region behind the sella and in front of the basioccipital.
Even the anterior and posterior parts of the sella are lined
with upgrowths from the parasphenoid.
The condition in the bird is paralleled by that of the reptile.
I n many vertebrates except the mammals (and some actin-
opterygian fishes?) the floor of the sella is covered by the
parasphenoid, yet it is this center of ossification in the croco-
dile which Parker (1880-1885, pl. 67, figs. VI, VII) identifies
as the basisphenoid. The lizard has thin trabeculae covered
below by a broad membral ossification which can only be
considered as parasphenoid. As in the bird, only the peri-
chondral and endochondral ossifications can be considered
truly basisphenoid.
The relationships of the parasphenoid to the basipterygoid
processes must be considered. I n the reptile and bird (T. J.
404 MALCOLM T. J O U I E
Parker, 1891, pl. 10, fig. 106; pl. 11, figs. 124, 136) there are
cartilaginous extensions from the trabeculae (basitrabecular
processes) which jut out as the anlage of the basipterygoid
processes. The fate of these cartilages appears to be similar
to that of the trabeculae - they are enclosed by the para-
sphenoid so that their own ossification is concealed. It is
quite probable that reduction in the size of the parasphenoid
has resulted in exposure of these bony processes of the basi-
sphenoid (many reptiles).
A somewhat different fate awaits the trabecula communis
in front of the sella. It disappears and its place is taken by
secondary ossification extending inward and upward from
the rostral part of the parasphenoid. This Ossification cannot
be considered as “presphenoid” or “orbitosphenoid” since
it does not correspond either in its relationships or manner
of origin with those centers of osszcation of the mammal.
Admittedly the above is an interpretation of observations
which have been viewed differently by previous writers. On
the basis of what occurs in the mammal (the source of our
bone names) this seems to be the best approach and is not
so different from the conclusions of Parker (1890).
Parasphenoid (figs. 1, 3-6, 17-19)

The parasphenoid arises from 7 membral ossification cen-
ters in the chicken (fig. 17) : an anterior rostral (rostropara-
sphenoid), a pair below the sella turcica (sellaparasphenoid),
a pair extending out from the dorsal margins of the sella
(alaparasphenoid), and a posterior pair of basitemporals or
.
basicranials (basiparasphenoid) It is possible that the num-
ber of centers varies in other birds. A multiple origin is in
agreement with the views of Jarvik ( ’48: 79; ’55) although
the number of centers differs.
The rostroparasphenoid arises as a median ossification early
in the 11th day of incubation (fig. 14). Within a day of its
first appearance its posterior end is fused with the basi-
sphenoid, somewhat later it fuses with the sellaparasphenoid.
HEAD SKELETOS O F CHICKEX 405
The juiictioii with the latter is detectable laterally and dorsally

f o r several days hy the dircctioii of the bony fibers. I n the
liawk and English Sparrow this junction is evident late in
the devclop~rieiitalperiod. A s a sccondary development the
rostroparasphenoid extends dorsally to either side of the
Fig. 14 l'l~oto~kiicrogrnpliof x-ncvtioii throogh tlic parasplienoid rostrum show-

ing the dcnnal bone and the cartilaginous iiiterorbitnl scptum. The joiiit betwecn
tlw roRtroD"r""piei1oicl n i i t l tlic. tlerinal ptrqgoicl is also shown.
tixl)cwilar communis and in sonip I)ii*tls,thus contrihutes to

the interorbital septum.
T1w sellaparasplienoid appears in thcl last half of the 11th
day of iiicubation as l d a t e r a l plates which soon fuse with
the liasispheiioid al)ovr aiid rostroparasplienoid anteriorly,
and somewhat l a t w with tlic basiparasphenoid plates (mid
12th (lay) posterioi-ly. Thcso plates scwd fihcrs of hone clor-
sally and medially and eventually form the floor of the sella
turcica (this floor forms from behind forward and from the
sides inward, closing occurs about the 16th day of incubation).
From this developing floor anterior and posterior upgrowths
line the sclla turcica (13th day). Posteriorly and laterally
Fig. 15 Photomic,rograph of x-section through the cartilaginous trabccula of

one side with its cnclosiiig basisphenoid adage of periclioiidral bone. The dermal
alaparasphenoid is seen extending out to the left.
the sellaparasphenoids extend down behind the eustachian

tubes and fuse with the basiparasphenoids (fig. 16). Fibers
f roni these origins eventually enclose the internal carotid
arteries to form the carotid canals.
The alaparasphenoids (= tympanic wings of Erdmann, '40:
361 ) are lateropostcrior membral processes which grow out
to citlicr side from the dorsal outer margin of the sella (fig.
IIEAD SKELETON OF CHICKEN 407
15). Each of these wings eventually forms the anterior and

dorsal walls enclosing the rostra1 air sac from the middle
ear cavity. Its dorsal surface forms a part of the floor of the
cranial cavity while the anterior plate extends down in front
of the custachian tube (15th clay of incubation). The latcral
Fig. 16 Photomicrograph of x-section through tlie area of fusion of the se1l:i-

a i d bxsiparasphcnoids bdiiud tlie custaehian tuhc. Tha ventral margin of n
triilirc.ul:i is seen above without evidence of nil iiivesting basisphenoid.
part of this anterior surface devclops first and soon fuses

wit11 the ventral plate of the hasiparasphenoid. The anterior
foranicn of the carotid canal is formed by failure of these
mai.giiis to close medially. If the basisphenoid ossification
of tlic basiptcrygoid process of the bird is exposed it would
be lwtween the margins of this plate and that of tlic sella-
pai=asplicnoi~l.
408 MALCOLM T. JOLLIF
The basiparasphenoid centers (= tympanic bones of mam-

mals), soon after their appearance (early 12th day of incu-
bation), fuse to the sellaparasphenoids. These broad plates
underlie the tips of the lagenae and spread out to cover the
base of the cranium. They later fuse along the midline and
with the deeper extensions of the sellaparasphenoids behind
the eustachian tubes. Soon after the appearance of these
plates (late 12th day) the anterior margin is extended for-
ward below the custachian tubes (beginning laterally), so
FASISPHENOID
SELLA~RASRIENOID
BASIWAWHENOID
LAGENAR omLm
OCCIPITAL CONDYLE
EXOCCIPITAL
-FORAMEN MAGNUM
Fig. 1 7 Ventral view of base of craiiiuin of chick at 12th clay of iiicubntioii

showing ~oiiieof the aiilageii of the pnrasphciioid, and the anlagen of tho basi-
sphenoid, aiid the exoccipitals.
that thcsc tubes are enclosed liclow by the 14th clay of incn-
bation ; the rriargins of the basiparasphenoid and the alapara-
sphenoid meet laterally in front of the eustachian tubes on
the 15th day. Laterally the basiparasphenoid forms thc ven-
tral margin of the tympanic cavity. Later in clevclopment
the basiparasphenoid comes in broad contact with the basi-
sphenoid through bony trabeculae (fig. 5 ) .
Basisphenoid (figs. 1, 3, 5 , 6 , 17-19)
The basisphenoid arises from a pair of perichondral cen-

ters which appear early in the 11th day of incubation (fig. 19).
This pair arises around the trabecnlar cartilages lateral to
HEAD SKELETOX OF CHICKEX 4-09
the posterior end of the rostroparasphenoid ; later these

centers fuse ancl encircle the sella turcica. Soon after these
centers appear they fuse to the posterior end of the rostro-
parasphenoid aiid somewhat later with the sellaparasphenoid.
Almost from the moment of its appearance the alaparasphe-
Iioid is fused to the dorsal margin of this bone. The para-
splieiioid eventually encloses the hasisplienoid laterally and
ventrally and even sends processes up into the sella which
form parts of the sellar tubcmlc arid the dorsum sellae.
At the vertical of the anterior end of the hypophysis, a
small perichondral process extends posteriorly from the ven-
tral margin of either trabccula. From this proiig a membran-
ous extension of b o n ~extends hack to the vertical of the
posterior wall of the sella wherc the bonc becomes perichon-
dral again. The dorsum sellac fornis in part about the anterior
tip of the notochord which lies just posterior to the sella
between the slightly separated parabasal cartilages. From
the ventral splints of the basispheiioid, bony trabeculae ex-
tend out aiid downward to uriite with the sellaparasphenoid
(12th day of incubation) and later with the basiparasphenoids.
Thr basisphenoid of the chickrii does not have basipterygoitl
processes although late (60 days of age) in the development
of the skull articular pads for tlic anterior ends of the pos-
teropterpgoids (see p. 420) dcvclop at the posterior end of
the ~.ostroparasplieiioid. Hasiptcrygoicl processes do develop
in niaiiy birds as outgrowth at the vertical of the sella
(hawks) ; such processes clewlop carly, as cartilaginous pro-
jections, before the time of hatching; later they appear to
he eiiclosed by the parasphcnoicl.
Orbitosyhenoid (figs. 1, 2, 5, 6 , 12)

The orbitosphenoid lias frequently been iclentified as the
latcro- o r pleurosphenoid, a new element found in some rep-
tiles and birds (Gooelrich, '30 : 392-393). This confusion has
bccn due to the introduction of the alisphciioid in the niam-
ma1 skull, changes in relationships between the different orbit
410 MALCOLM T. J O L L I B
bones due t o expansion of the orbit, or brain case, or both,

and to misunderstanding of the number of bony units in this
area (Goodrich assumed that both a “ pleurosphenoid ’ ’ and
a n orbitosphenoid occur in birds whereas actually only one
element is present).
The orbitosphenoid is a perichondral development which
arises from two centers. It appears during the 14th day of
incubation. The dorsal center is triangular, the apex directed
down ; the ventral center is irregular but forms a more-or-less
transverse plate above the alaparasphenoid margin. It is
quite probably that these centers represent the “orbito-” and
“pre-” sphenoid centers of the mammal. There is a large
fontanelle between the two parts which is restricted as ossi-
fication proceeds until it finally closes (26 or more days after
hatching).
The orbitosphenoid lies in the posterior wall of the orbit,,
helow and slightly overlapped by the orbital extension of
the frontal, anterior to and partly overlapped by the squamosal
and abutting ventrally on the dorsal margin of the alapara-
sphenoid. The second and third branches of the trigeminal
nerve pass through a large foramen between the postero-
ventral margin of this bone and the parasphenoid (see
Icesteven, ’25). The optic nerve emerges medial and anterior
to this plate while the oculomotor (111) and trochlear (IV)
penetrates it lateroposterior to its optic nerve margin. The
orbitosphenoid forms the entire postorbital process in the
chicken, the Great-horned owl (Bubo virginialzzcs [ Gmelin] ),
and the English Sparrow; whereas in the hawks examined,
Mallard and pigeon the lateral aspect of this process is
covered partly by the squamosal. The interrelationships of
the various bones at the postorbital process is variable in
rliffcrmt species.
8qzcninosal (figs. 1, 2, 4-6, 18, 19)
The squamosal is a membral ossification which appears late

in the 10th day of incubation at the quadrate articulation.
HEAD SKELETON OF CHICKEX 411
When fully devclopetl it covei*sthe latcval aspect of the brain

case. hhove, it ovci*lapsthe frontal and parietal; behind, it lies
i l l contact with the supra- aiid exoccipitals. Anteriorly, it
overlaps the orbitospheiioid ; the tympanic cavity and the
quadrate lie bclow it. The squaniosal lias an articular socket
for the outer head of the quadrate and a short “zygomatic”
process which juts down along the postclrolatcral margin of
the quadrate. A large air space from the middle ear cavity lies
iiiside tlic posteroventral margin, posterolateral to tlie dorsal
margin of the prootic.
STROPARASPHENOI
I i
Fig. IS Aiitrrolntrrnl view of skiill and ninndible of thc chick at the 11th day
of incubation.
Rittiwr ( ‘1% - sec Owm, 1849, pl. 39, figs. 1-3; 1879: 120,
pl. XSXI, figs. 1-3) described a small dermal ossification on
the postorbital process of birds which he relates to the
postorbital bone of the reptile. Further (p. 17) he states “Bci
allen untersuchten Vogelschadcln ohm Ausnahmc war es
rnir rnoglich, das Postfrontale nachzuweisen, . . .” This state-
irient is followed by a lengthy list of species supposedly having
a postorbital. What Bittner observed in the species for which
he had developing skulls, was a secondary membral ossification
covering the postorbital procrss.
I n the gallinaceous birds, not only the postorbital but also

the zygomatic process of the squamosal are extended by such
ossifications which fuse distally to form a closed temporal arch
resembling that of a reptile. Ossifications of this nature also
occur in the formation of sesamoids, or bony ligaments. If
the bone that Rittncr describcd as the postorbital were a true
member of the cranial corriplex it would appear a t approx-
imately the same time as the other orhit-rimming bones and
would lie between adjacent bones rather than being an os-
sification in the postorbital ligament overlying whatcver boiic
is involvetl in the postorbital proccss (orhitosphcnoicl, frontal,
parietal, o r squamosal in diff ercnt birds). Parker (1869 : 790,
P1. 84, figs. 13,14) clescrilied a postorhital in a three week old
chicken: The bone which he figures is a part of the or-
bitosphenoid. The fact that he observed it oiily in this oiic
instance suggests it is an aiiorrialy and riot propcrly a postor-
bital. Lack of a postorhital is n good distinguishing charactw
for the Class Avcs (hut would thus exclude Archncopteryr).
I’nrietcrl (figs. I , 2, 4 , 5 , 7,8, 12)

The parietals arc niemhral bones appearing early in the
12th clay of incubation, but they are not distinct until early
in the 14th day. From this time on they spread out rapidly
over the postcrodorsal aspect of the cranium, particularly
forward. The aiilage appears along the ridge between the
cerebral and cerebellar fossac and in the definitive plate the
bone is much thickened here to form an inner ridge. Thc
parietal contact tho frontal along its clorsoanterior margiu,
is overlapped by the squarnosal latc~rally,and contacts the
supraoccipital ventroposteriorly. The two parietals meet at
the midline along a fairly straight suture line. A tlis7erticulum
from the middle ear cavity penetrates the posterolnteral angle.
Int erpci rie t m 1

Heilmann (’26, fig. 3) has illustrated the intcrparietal in
Anser anser Linnacus. In thc chicken there is no eviclcrice,
at any stage, of its presence in the cranium. Staurenghy (see

Strong’s “Bibliography of Birds” for references) wrote
several papers on the occurrence of this bone in birds but I
have not seen any of them.
Frontal (figs. 1, 2, 4,5,12,18)

The frontal arises early in the 11th day as a thin line of
membral ossification along the supraorbital margin. Erdmann
( ’40: 336) recognized two centers for this bone, the posterior
of these he equated with the postfrontal. If two centers
appear they must be considered parts of the frontal. From
this origin it gradually spreads medially, posteriorly and down
into the orbit; the last area to be covered is the midline just
above and in front of the parietals where the plates of either
side eventually meet along a somewhat irregular suture. The
anterior tip of the frontal is pointed and bent laterally to
expose the mesethmoid (perpendicular plate). The anterior
tip underlies the posterior tip of the nasal. Posteriorly the
frontal contacts the parietal ;laterally the squamosal overlaps
it while in the orbit it overlaps the orbitosphenoid. The suture
between the frontal and the parietal is largely a simple
contact but near the midline the frontal slightly laps the
parietal. The frontal sutures are the last to close in the
cranium proper.
Nasal (figs. 1,2,5,18)

The nasal, a membral bone, appears early in the 11th day
of incubation along the posterior margin of the narial ap-
perture. The medial process appears about a day later and
gradually extends forward (16th day). The nasal contacts
the premaxilla with its dorsonarial process and the premaxilla
and maxilla with its postnarial limb; posteriorly it overlaps
the anterior ends of the frontals and contacts the prefrontal
along its lateroposterior margin. I n the hawks, the nasal
fuses with the secondarily ossified vestibular wall.
414 MALCOLM T. J O L L I E
Prefroatal (figs. 1,2,5,18)

The prefrontal is a membrane bone which in the chicken
appears early in the 11th day of incubation and has one or
two centers of ossification (fig. 12; Erdmann, ’40: 335) ; the
orbital process may ossify separately but soon fuses with
the main supraorbital part. The prefrontal abuts against
the lateral aspect of the nasal and the anterior end of the
frontal. The ventral tip of its orbital process is connected
with the labial bar by a ligamentous tie. The English Sparrow,
House Finch and Junco lack the prefrontal and only a vestige
remains in the Robin (see figures of Parker, 1874-79, for
other passerines).
The prefrontal is usually identified as the lacrimal probably
because of the similarity of its position to the lacrimal of the
mammal (Cuvier, 1835, 11: 579-580: Owen, 1879: 164-165).
Since the bird is related to the mammal through the reptile,
any direct comparisons should be made with the latter group.
I n the reptiles both a prefrontal and a lacrimal are usually
present although the lacrimal may be missing in some. The
lacrimal lies in front of and lateral to the lacrimal duct but
its inner surface may be grooved or even enclose the duct
(crocodile). The prefrontal has a supraorbital division above
the level of the lacrimal bone which forms a part of the outer
surface of the skull and an orbital process which passes
downward medial to the lacrimal duct. The orbital process
forms a part, or all of the bony anterior wall of the orbit.
I n the bird the bone in question is formed much like the
prcfrontal of the crocodile o r lizard and has the same relation-
ship to the lacrimal duct. The exceptions to this statement
are ratites. Dromaeus,Rhea and Apteryx have the “orbital”
process perforated for the passage of the duct. There is no
evidence that the outer splint is ever a separate lacrimal bone
but its relationships are suggestive of this.
The assumption (Erdmann, ’40 : 335) that this bone in the
bird is equivalent to a fused prefrontal (supraorbital part)
and lacrimal (orbital process) is not supported by the posi-
tion of the later in relation to the lacrimal duct (see also

Marinelli, ’36: 819; Gregory, ’20: 234). Changing the name
of this bone seems proper in view of the above. Stresemann
(’27-36: 52, 53) has already done so.
Nesethmoid (Perpendicular Plate of Ethmoid -

figs. 1, 2, 5 )
The mesethmoid is the anterior portion of the sphenethmoid
plate of the primitive amphibian and early reptile (the orbito-
sphenoids are the posterior part). The mesethmoid appears
during the 17th day of incubation (a chondromembral os-
sification) and gradually expands to the adult dimensions
when along with the secondary ossifications in the interorbital
septum, it forms an interorbital plate.
I n some species (hawk), the interiiarial septum is ossified
(fuses with ossification in the vestibular walls). This sec-
ondary ossification is separated from the mesethmoid by the
craniofacial fissure which allows movement of the rostrum
along the frontonasal hinge. I n the Ostrich, the septa1 and
mesethmoid ossifications are continuous (the mesethmoid os-
sifies well before the nasal septum). The nasal capsule and
septum may ossify as a unit (falcons) or small ossicles may
develop at different points in the capsule. Parker (1875-79 a
and b) has called some of these ossifications “septomaxil-
laries” and “medial maxillaries. ”
The mesethmoid of the chicken is exposed dorsally between
the anterior tips of the frontals and the posterior tips of the
nasal processes of the premaxillae. I n most birds there is
very little of this bone exposed here, but in ratites, swifts and
some gallinaceous birds there is a noticeable exposure.
Lnteral ethmoid
A lateral ethmoid is represented in the reptiles and mammals
only by a small process of the chondrocranium (pars plana).
Ossification is usually present in the bony fishes. It is a
secondary chondral membral ossification which appears at the

very end of the developmental period (hawk, owl, passerine)
and as such indicates its phylogenetic newness in the bird.
This plate may form a part, or the whole, of the anterior wall
of the orbit. Laterally this ethmoid (antorbital process) may
contact or fuse with the orbital process of the prefrontal.
Richardson ('42: 330) has confused this plate with the pre-
frontal, although both are present in Sitfcr.
UGENAR OTOLITH&+
NOTOCHO~O\
SACCULAR, UTRiCULAR OTOLIT6

Fig. 19 Ventral view of skull of the chick at the 11th day of incubation.
Preinaxilla (figs. 1-3, 5,18,19)

The premaxillae appear early in the 11th day of incubation
as bilateral membral ossifications which gradually meet at the
midline and fuse (16th day). The nasal processes (or frontal
processes) extend back gradually and eventually overlap the
anterior end of the mesethmoid. The nasal processes remain
as bilateral structures throughout most of their lengths, only
the extreme anterior ends are fused together at the midline
(as are the bodies of the premaxillae). The labial process

extends back to the vertical of the frontonasal hinge. The
palatine process appears last-early in the 12th day of
ASIBRANCHIAL I
PREARTICULAR
Fig. 20 Ventral view of mandible and hyoid apparatus of the chick a t the
11th day of incubation.
Fig. 21 Palate of nestling Robin about 10 days of age.

Fig. 22 Palate of adult Robin with anomalous relation of vomer and palatine
on the left side (reader’s right).
Fig. 23 Lateral view of palatine-vomer junction in nestling Robin.
Fig. 24 Lateral view of palatine-vomer junction in adult Robin.
incubation. It is a posteriorly projecting sheet, pointed at

its tip, which overlaps (as viewed ventrally) the anterior
end of the palatine but does not fuse with it.
I n passerines the palatine process (called the palato-maxil-
lary in recent papers on f ringillids -Amadon, ’50 ; Tordoff,
’54 - see figures of Parker, 1877-79, and pp. 252-253) is long
and thin in the nestling with a constriction at its base. In the
Robin (figs. 21-24) or English Sparrow it eventually fuses
to the anterior end of the palatine and is indistinguishable
from that bone. I n the Junco this process appears in the
nestling but remains free of the lateral margin of the anterior
end of the palatine. Later its basal constriction deossifies
(as a result of hinge movement in this region?) so that a free
“palato-maxillary ” results. I n other birds the palatine pro-
cess may fuse with the labial process or with the maxilla.
& f a d l a (figs. 1-3,5,18,19)

The maxilla is a membral bone that appears early in the
11th day of incubation. It is reduced in size and contributes
little to the margin of the upper jaw but forms a part of the
surface of the palate, lateral to the anterior end of the palatine.
Medially it lies dorsal to the anterior end of the palatine
and sends a small, sheet-like, palatine process toward the
midline. I n the chicken these processes lie anterior to the
internal median naris while in the passerines they lie lateral
to this opening. Laterally the maxilla has a slim labial process
which extends posteriorly along the line of the upper lip. I n
many birds there is a dorsally projecting postnarial process.
The palatine process is much inflated in some birds and may
largely fill the base of the bill. I n palaeognatlis this bone
forms a much larger part of the palate than in most neognaths.
Jugal (Malar or Zygomatic Bone of Hu,rnan-

figs. 1-3,18,19)
The jugal is a small splint of membral bone which appears
early in the 11th day of incubation. It lies above the labial
process of the maxilla and the anterior end of the quadrato-

jugal and forms the middle part of the labial arch (the
zygomatic, suborbital, or quadratojugal arch of some). A
jugal is lacking in the Magpie, English Sparrow, Robin and
Kestrel (Falco tiuLlzuncuZu,s - Sushkin, '05 : 3 6 ) .
Quadratojugnl (figs. 14,12,18, 19)

The membral quadratojugal is the first bone of the skull to
ossify, being indicated late in the 10th day of incubation. It
forms the posterior part of the bar which margins the upper
lip (labial arch) ; its posterior articulation with the quadrate
is of the ball and socket type.
Quadrate (figs. 1-4, 6, 12, 18, 19)

The quadrate appears early in the 11th day of incubation
as a perichondral ossification. Ossification continues until
the chick is fully grown. The quadrate has proximally two
articular facets, separated only by a line. The outer facet ar-
ticulates with the squamosal, the inner with the prootic. I n
the ratites these two facets arc well separated but not strongly
outlined. They are distinct in the tinamou but close together
(separated by a notch), as in many species (including the
chicken). Separation of these heads is due to penetration
by a diverticulum from the inner ear cavity between the
squamosal and the prootic (a second diverticulum enters the
body of the quadrate). The body of the quadrate has a n
articular area for the pterygoid on its anterior inner aspect.
This articular area extends up onto the ventral margin of the
broad orbital process which projects anteromedially.
Pterygoid (figs. 1, 3, 5, 6, 18, 19)

The pterygoid of the bird can be developed as a single
unit (Palaeognathae and some Neognathae) or in two parts
with an articulation between them. This condition has been
described by Parker (1872 : 224 ; and subsequent works) and
unequivocally stated by Pycraft (1898 : lix ; ’00-’03 : 349) and

Kingsley ( ’25: 180). The need for an exacting terminology
for this situation in most neognathine birds has not been met.
For the sake of simplicity and clarity the two parts can be
called the anteropterygoid (mesopterygoid or hemipterpgoid)
and posteropterygoid (usually identified as the pterygoid) .
The pterygoid of A p t e r y x is unique in that it is forked
anteriorly : one splint, representing the anteropterygoid, con-
tacts the vomer while the other splint, passes forward above
the ‘‘lateral palatine” process of the maxilla. The lateral
splint resembles in its position the transpalatine (ectoptery-
goid) bone but it arises as an integral part of the pterygoid
according to T. J. Parker (1891, pl. 11, fig. 121 ; pl. 8, fig. 67).
The antero- and posteropterygoids ossify separately in the
chicken but share a common mesenchymatous anlage (fig. 19).
Both appear early in the 11th day of incubation. The anterior
part, from the time of its appearance, is fused to the palatine;
it is slightly differentiated by the direction of its fibers and
their slightly heavier ossification. This anterior portion lies
along the parasphenoid rostrum and forms the “pterygoid
articulation” of the definitive palatine bone. The anterior end
of this part of the pterygoid does not extend forward to the
vomer, rather a rostra1 process of the palatine (see under
palatine) extends forward from its tip to the posterior end
of the vomer. Pycraft ( ’00-’03 : 350) has described the antero-
pterygoid as a peg like anterior projection of the postero-
pterygoid in the “Galli.” This is the case in the “Anseres”
but not in the galliforms.
The posteropterygoid appears slightly earlier than the
palatine and anterior part. It lies at an angle in the roof
of the mouth, extending from the posterior end of the palatine
to the quadrate. The anteromedial end is irregular at first and
a distinct articulation with the anteropterygoid does not
develop until shortly after hatching, The medial aspect of the
anterior end of the posteropterygoid clasps the parasphenoid
rostrum and eventually develops an articular surface which
glides over a similar surface on the rostroparasphenoid. The

posterolateral articulation with the quadrate develops late ;
it is complex in form and has contact both with the body and
the orbital process of the quadrate.
In the hawk, pigeon, Robin and other passerincs examined,
the pterygoid arises as a single unit which may contact
anteriorly the vomer (no vomer in pigeon). Late in develop-
ment the anterior end fuses to the palatine and an articulation
develops between it and the posterior part (figs. 23, 24).
Amadon ( ’50: 217) noted that in some oscines an articulation
does not develop even though the anterior part fuses to thc
palatine. I n some birds the pterygoid is jointed but the
anterior part is not completely fused to the palatine. Some
bustards (Neotis cnfra denhnvni [Children], American Mu-
seum 1967) have partially separate anteropterygoids which
articulate with the posteroptcrygoids and contact the vomer
between the palatine.
Palatine (figs. 1,3, 5, 6,18,19)

From the above it is apparent that we also need to redefine
the palatine. In those species having a unit pterygoid there
is a true palatine, whereas in most birds the anterior part of
the pterygoid is fused to the palatine to form a compound
bone. This compound bone can bc called a pterygopalatine.
The form of the palatine, or the pterygopalatine, is deter-
mined by the attachment of the palatine muscles and is
extremely variable. The terminology applied to this structure
(Amadon, ’50 ; Tordoff, ’54 - taken from Parker, 1877-79)
is hardly appropriate. The “prepalatine process” is the an-
terior end of the palatine and not a process in the strict
sense. The “transpalatine process” is described by Parker
(p. 109) as arising as a separate ossification (Chelidorz urbica,
P1. LII, fig. 4). This is the case generally in the aegithog-
nathous types, where it appears very late in development and
almost immediately fuses with the palatine. It can only be
considered as a specialization, an the basis of its origin and
since, in lower birds in general, any process that appears

arises as a ventrolateral extension of the posterior end of
the palatine. This process cannot be considered as having any
relationship to the transpalatine bone (ectopterygoid) of the
reptile ;theref ore, it is misleadingly named, whether homology
or disposition is considered. It might better be called the
posterolateral process. The “interpalatine process ” (includ-
ing the medio-palatine process) might better be called the
choanal process since it margins the median slit of the
internal naris. The postpalatine process in some birds is fused
with the anteropterygoid. I n the adult duck or swan, or the
nestling of many species, it is free and may develop an
articulation with the anterior end of the postpterygoid.
Another process, observed in the young and indicated in
some adults, can be called the rostral process (Parker’s
ethmopalatine process). I t extends forward along the para-
sphenoid rostrum (or anteriorly along the line of the rostrum),
above the posterior end of the vomer. Later it may fuse to
the posterior end of the vomer, as in the oscines (figs. 21-24).
I n the chicken a pterygopalatine appears early in the 11th
day of incubation. The membral palatine portion is a twisted
strap extending from the anterior end of the posteropterygoid
to the anterior end of the maxilla. The rostral process soon
develops and its anterior end gradually extends to the posi-
tion of the posterior tip of the vomer (early 13th day of
incubation). The anteropterygoid is fused to the palatine
from its first appearance and forms the pterygoid articular
process. There are no posterolateral or choanal processes.
The rostral process is small and pointed. The anterior tip of
the palatine is gripped between the palatal process of the
premaxilla and the anterior end of the maxilla. I n many
oscines the anterior end of the palatine fuses with the palatine
process of the premaxilla and this fused band may develop
a transverse hinge where it contacts the palatal body of the
premaxilla (figs. 21, 22). I n other birds the palatine may fuse
to the body of the premaxilla, the palatine process remaining
free or fusing with the labial process or maxilla.
HEAD SKELETOPI' O F CHICKEN 423
Vonzer (figs. 1, 3, 5, 6)
The membral vomer arises early in the 13th day of incuba-
tion as bilateral splints of bone (Parrington and TVestoll, '41,
have clearly demonstrated the homology of the vomer of the
mammal with that of the reptile or bird). Erdmann ( '40 : 333)
states that only a single center occurs in the chicken. Such an
interpretation is plausible since with ossification the anterior
parts of the two centers fuse. Occasionally the double origin
is detectable in whole specimens ; cross-sections more clearly
reveal it.
The splints forming the vomer lie in the internarial septum
and almost immediately the anterior halves, or more, fuse
across the midline. The posterior splints remain separate and
lie below the anterior tips of the rostra1 processes of the
palatines. The anterior tip of the fused vomers lies between
the posterior tips of the palatine processes of the maxilla.
I n nearly grown chickens (70-90 days) the vomer may disap-
pear as an ossification.
There is a wide range of variation in the relationship of the
vomer to the other bones of the palate and its size and shape ;
it may be lacking as an ossification (cathartid vultures,
pigeons) or involve an endochondral invasion of the nasal
cartilage (Parker, 1872 : 224 ; 1875-79 : 108-109).
Sclerotic eye ri9tg (fig. 1)

The number and arrangement of sclerotic plates is variable
among individuals and in either eye (see Lemmrich, '31;
Curtis and Miller, '38). A count of 6 rings showed numbers
ranging from 11-15. There are generally 14 which is in agree-
ment with Steiner's ('52: 372) statement, however 16 seems
to be the basic number of plates in a pelican (4 rings counted),
and 11 in the pigeon (6 rings counted). The plates first
appear during the 12th day of incubation and are distinct by
the 13th day. The plates are broader posteriorly, broadest
in the posterodorsal quartile of the ring. The plates overlap
one another in a seemingly irregular manner.
ACCESSORY BONES
Several small secondary ossifications are associated with

the skull in different birds, none of which occur in the chicken.
Supraciliary
Ossifications within the supraorbital membrane occur in
several species. I n the tinamou the entire area may be filled
in with an irregular series of plates while in the accipitrine
hawk a single plate, which articulates with the tip of the
supraorbital process of the prefrontal, is present usually. As
a secondary ossification, the supraciliary (or supraciliaries)
appears late.
Ossiculuin lacrivnopalatinuin
The lacrimopalatinum (Brandt, 1839 : 4 ; Beddard, 1898 :
136) is present in such diverse types as Diomedea, Morus,
Cariarna, Turacus and Eudynamis. I t is usually confused
with the ossiculum suprajugale and the lacrimosuborbitale.
The lacrimopalatinum lies between the tip of the orbital
process of the prefrontal and the lateral margin of the palatine
directly below. In effect it binds the orbital process to the
labial bar and to the margin of the palatine. I n shape it
is broad at its articulation with the prefrontal and tapers
to its distal tip.
Ossiculum suprajugale
An ossiculum suprajugale (Brandt, 1839: 2) occurs in
cormorants and anhingas as a large ossicle lying above the
labial process of the maxilla.
Ossiculzm lacrimosuborbitale
Nitesch (1811 : 76) described a small accessory bone found
in the terns of the Genus Sterna (observed also in a specimen
of Larus nrgeNtatus-also figured [?I for Catharacta by
Maillard, '48, fig. 34A, e). This ossicle extends from the
outer margin of the ventral tip of the orbital process of the
prefrontal posteriorly along a line below the eye. Since this

accessory has not been formally named (called accessory
bone by Nitzsch) I have given it the above name.
Ossicula articularia
Magnus (1871 : 100) described as ossa acccssoria two small
ossicles lying in the posterior wall of the articular capsule
of the quadrate-mandibular joint in crows, Coccothraustes,
Lanizcs, and Cuculus. These sesamoid ossicles are present in
most, if not all passerines, Capitonidae, Rhamphastidae and
probably in other families allied to the later. Only a trace
of the lateral ossicle was seen in a specimen of Colaptes and
a small posterior one was seen in J y m . Both were observed
in Bartramia. A large lateral ossicle was found in a specimen
of Merops; this ossicle was observed in Numenius, Limosa,
and Totanus. A posterior ossicle was observed in specimens
of Halcyon, Dacelo, Buceros, Lophoceros, Larus, and Pandion.
None was observed in other “aegithognathous” types such as
Apus, or Caprimulgus, or in forms close to these-Trogon,
Steatornis and pigeons.
Nitzsch (1811: 74-77) described as ‘‘ossa palatomaxillaria”
ossifications at the angle of the mouth in Fulica. They are
ossified insertion tendons of the adductor mandibulae muscle
and not properly accessory bones.
0 s opticurn
The 0s opticum is found in many different kinds of birds
(=os opticus of Tiemeier, ’39; ’50). It forms around the
optic nerve where it penetrates the sclera of the eye ball.
It may be a single crescentic piece or two lateral pieces. I n
a specimen of Colaptes cafer (Gmelin) there is a horse-shoe
shaped ossicle about the optic nerve and a second ovoid os-
sification lies well above the optic nerve. Tiemeier ( ’39 : 334)
found this ossicle in Gallus, but my oldest stages do not show
it.
0 s sip honnium
The 0s siphonium appears in many passerines (Strese-
mann, ’27-34: 60, fig. 64-after Nitzsch, 1811: 30, Taf. I,
figs. l”, 1 +) and also in non-passerines such as A p u s npus
(Linnaeus) and Colnptes ccl.fer. It is a secondary ossification
in the walls of the diverticulum of the middle ear cavity which
in the Raven (Corvus coraa) extends down to the mandible.
This tubular ossification passes through a notch in the tym-
panic margin. I n the flicker only a small nodule of bone
occurs in the tube.
0 s mchale
A membrane bone occurs in cormorants and anhingas in
the nape region. This is a long spine which projects poste-
riorly from its articulation with the mid-dorsal point of the
nuchal ridge of the cranium. It serves as an area of origin for
a specialized (separate) part of the complexus muscle. I
have given it the above name (see Beddard, 1898: 411, foot-
note).
MANDIBLE
The bones of the lower jaw are usually 6 in number, and

include all those known for the reptile with the exception of
the coronoid. Parker (1890: 14) noted that “The coronoid
does turn up in the Passeres, and in some Grallae,” but, in
his description of the crow (1872: 222) he has called the
prearticular the coronoid. The only evidence I have of a pos-
sible coronoid is in the mandible of a Golden Eagle at the time
of hatching (see below).
Articular (figs. 1,5)

The articular appears during the 17th day of incubation as
an ectochondral ossification enclosing the dorsal, anterior and
part of the ventral aspects of the tip of the medial articular
process of hleckel’s cartilage. This ossification is partly fused
from its very beginning with the tip of the articular process
of the prearticular. It is not fully developed in chickens until

full growth is achieved (about 100 days) when it covers
the entire articular area for the quadrate and has a strong,
upcurved, retroarticular process. It does not fuse completely
with the angular and supraangular until the period of general
fusion of skull bones (90 days or more). I n the English
Sparrow the articular remains free of the fused posterior ends
of the angular and prearticular until the time of nest-leaving.
Angular (figs. 1,5,18,20)

The angular is a membral bone which appears late in the
10th day of incubation as a splint lying along the ventral
margin of the mandible. It has a retroarticular process and
contacts the prearticular medially. Anteriorly it tapers to a
point between the dentary and the splenial plates. I n the
English Sparrow the angular is large and replaces much of
the ventral marginal portion of the prearticular.
Supraamgular (figs. 1,5,18,20)

The membral supraangular appears late in the 10th day of
incubation and, like the angular, it has a retroarticular process.
The processes of these two bones extend posteriorly, along
with the retroarticular process of the articular; their tips fuse
at about 75 days of age. By 90 days these membral bones
and the endochondral articular are fused into a single unit.
The supraangular covers the lateral aspect of the proximal
end of the mandible ; contacting the dentary along its dorsal
margin and the angular along its ventral margin. The (dor-
sal) marginal process of this bone comes to lie between the
outer lateral plate and the inner, marginal process of the
dentary. I n many birds a fenestra penetrates the mandible
between the margins of the supraangular and the dentary,
this is the anterior mandibular fenestra. I n the passerines,
and many other birds, the supraangular has a large fenestra
in it, the posterior mandibular fenestra.
Dentary (Dentosplemial- figs. 1,5,18,20)

The dentary arises from two membral centers that appear
early in the 11th day of incubation (The dentale and mento-
mandibulare of Erdmann, '40: 368-369, Abb. 1). One center
forms the anterior end of the ramus and has a posterior
dorsal marginal process (mentomandibulare) , the other arises
along the outer ventral margin and sends a plate of fibers
up over the lateral aspect (dentale of Erdmann = anterior
splenial of primitive amphibian - see Romer, '49, fig. 146).
By the 13th day of incubation the anterior tip of the postero-
lateral plate begins to fuse with the anterior part and fusion
soon makes it difficult to delimit the two. The lateral plate
extends dorsally to the margin of the ramus where it lies
outside of the anterior tip of the supraangular ; the marginal
process of the anterior center lies inside this anterior tip.
Almost from the moment of their first appearance, the an-
terior centers are partially fused across the midline. Later
a symphyseal ridges rises upward from the joint, behind the
anterior ends of the Meckel's cartilages. This ridge is ob-
literated with the covering over of the cartilages by later
ossification.
Prearticular (Complenzemtare or gomiale -

figs. 5, 18, 20)
The membral prearticular appears early in the 11th day
of incubation but does not become distinct for several days.
It has a broad median articular process and a short anteriorly
projecting body which lies on the medial aspect of the man-
dible. This bone extends forwards for about one eight of
the length of the mandible. I n chicks 34 days of age it is
fused with the articular but not to the rest of the mandible.
It gradually develops a retroarticular process (65 days of
age) which becomes indistinguishably fused with this part
of the articular at about 75 days of age.
DeBeer ('37: 281) and Marinelli ('36: 829) state that a
prearticular is lacking in Callus. This error can probably be
traced back to a similar statement by Parker (1869: 790).

Marinelli, following Stresemann ( '27-34 : 58), lists it as mis-
sing not only in the chicken but also the sphenici, Struthio,
Columbae and Striges. Erdmann ( '40: 374) was the first to
record this bone for the chicken. A prearticular is apparently
present in all birds although it varies considerably in size.
The prearticular of the chicken differs from that of most
birds in that it does not extend far forward. I n some birds
it extends for nearly one half the length of the ramus. The
anterior end of this bone may be a conspicuos feature of
the internal aspect of the mandible (Procellarius, Morus,
Nycticorarx;). I n the English Sparrow it is much reduced;
posteriorly it fuses with the angulare at an early stage
while anteriorly it is a thin irregular splint of bone. I n both
the sparrow and the Robin the posterior subarticular ex-
pansion of this bone is much less than in the chicken.
Sple.lzial (figs. 5, 18, 20)

The membral splenial (= posterior part of splenial -see
Romer, '49, fig. 146) appears early in the 11th day of incuba-
tion as a splint lying just inside of the ventral margin of the
mandible. A thin sheet of bone extends upwards from the
splint to cover much of the inner aspect of the mandible
throughout the middle half of its length. Posteriorly the
splint lies above the angular and extends back to the anterior
end of the prearticular. Anteriorly the tip of the splint fuses
with the dentary. The dorsal sheet remains free from the
dentary until full growth is achieved. In the chicken this
sheet covers the inner opening of the anterior mandibular
fenestra but in many birds this is not the case.
Coromid (fig. 25)

In the Golden Eagle, at the time of hatching, a small splint
of bone was found lying just anterior to the tip of the pre-
articular. Such a splint was present in either ramus. The
very regular outline belies its being a detached tip portion

of the prearticular.
BRANCIAL SKELETON
Colzcmella (figs. 1, 12, 13)

Ossification of the columella (= stapes, =hyomandibula
of some vertebrates) begins during the 15th day of incubation
and proceeds through the entire growth period to the mature
condition.
I * *
The hyoid apparatus of the bird is simple in form but the

parts are difficult to homologize with those of other ver-
tebrates. Because of this, the names used in Kewton and
?JJF'WANGULAU
SPLENIAL AN'GULAR
Fig. 25 Medial aspect of mandible of a Golden Eagle chick at the time of
hatching.
Gadow (1893-96 : 453 and figure) will be employed. I n paren-

theses are indicated the names which perhaps more accurately
indicate the homologs. The basal portion of the hyoid cornua
(basibranchial) begins to ossify early in the 11th day of
incubation; the other parts do not ossify until late in develop-
ment.
Entoglossal (Hypohyal -fig. 2 6 )
The entoglossals are bilateral cartilaginous and bony struc-
tures in the tongue. The anterior ends of these usually remain
cartilaginous and fuse across the midline. The entoglossals
articulate with the anterior end of the copula or basihyal.
Basihyal (Basibranchial I -fig. 26)

The basihyal (-hyal refering more to hyoid apparatus than
hyoid arch) is the central unit of the tongue skeleton. The
entoglossals articulate with its anterior end, the first basibran-

chials and the urohyal with its posterior end.
Basibranchial I (Hyopbranchial I -fig. 26)

The basibranchials are the first parts of the hyoid apparatus
to ossify-beginning in the 11th day of incubation. Ante-
riorly they articulate with the basihyal and posteriorly with
the ceratobranchials.
ENTOGLOSSAL
CE.QATCBRbNCHIAL
BPISIBRANCHIAL I
Fig. 26 Ventral view of incompletely ossified hyoid apparatus of chicken

3 months of age.
Cerat o bram9zial (fig. 26)

The ceratobranchial continues the first branchial arch up
from the basibranchial. The length of this unit varies greatly
in different species.
Urohyal (Basibranchial II -fig. 26)

The urohyal extends posteriorly from the end of the basi-
hyal. It lies on the underside of the thyroid cartilage and is
sheathed by the connective tissue covering that structure.
DISCUSSION
The head skeleton of the chicken appears to agree in most

details with that of other birds which differ from their reptile
progenitors by lacking postfrontal, postorbital and ectopter-
ygoid bones. The bird agrees with the reptile in the great
reduction or usual absence of the interparietal (postparietal).
The chicken has usually been considered a primitive bird
but such a conclusion is not supported by its head skeleton.
The very early separation of the anterior and posterior parts
of the pterygoid coupled with the reduction of the vomer and
the lack of basipterygoid processes suggests a marked diver-
gence from the ancestral type, at least as regards the skull.
This modification perhaps is adaptive in nature.
Lack of the lateral ethmoid ossifications might be conceived
as primitive, as might also the schizognathous palate, the
broad nature of the supraoccipital and the comparatively early
filling in with bone of the the entire occipital region. The
angular, narrow cranium is usually thought of as primitive
but there is no evidence to support such a view.
It is difficult to determine what is, or is not primitive; cer-
tainly the choice of such characters is going to be somewhat
arbitrary and can only be based upon a knowledge of the
comparative anatomy of birds. Detailed studies of the de-
velopment of the head skeletons of other birds are desirable.
SUMMARY
The time of appearance and the morphogenesis of the

different bones of the head of the chicken have been described
along with some comparative remarks on the same bones in
other birds. In the chicken, bone first appears late in the 10th
day of incubation. Ossification and expansion of bones con-
tinues until about 100 days of age at which time full growth is
achieved. The endochondral bones of the chicken appear
first as membral, perichondral ossifications with secondary
invasion and replacement of the cartilage. The interiors of
all of the bones are filled with trabeculae or are hollow. Air
sacs penetrate even amongst the trabeculae.
Fusion of the skull and mandibular bones bcgins in the oc-
cipital region at about 75 days of age and continues forward
and upward through the cranium until closure of the frontal
sutures, and most of the sutures of the mandible, at about
100 days of age. The prefrontal remains separate as do the
posterior tips of the nasal processes of the premaxilla, and
the ventroanterior end of the nasal and its medial process.
The sutures of the dentary and splenial are also retained
to a certain extent.
The bone usually called the lacrimal corresponds more
nearly to the reptilian prefrontal and has therefore been given
that name. The basisphenoid bone is very small and restricted
to pcri- and endochondral ossification of the trabeculae en-
closing the sella turcica. This bone cannot be seen from below
due to its investment by the parasphenoid. The parasphenoid
has 7 centers of ossification and forms most of the basis
cranii, largely enclosing the basisphenoid. The pterygoid is
divided into two parts from the beginning of ossification. The
anterior part fuses with the palatine and articulates with the
posterior part which is usually called the “pterygoid.” The
dcntary is found to be a compound bone comprising the
dentary and the anterior splenial. The bone usually called
the splenial is the posterior splenial. I n the Golden Eagle
a trace of the coronoid was found.
The bird differs from the reptile in lacking the postfrontal,
postorbital and ectopterygoid ; it agrees with the reptile in the
great reduction or usual loss of the interparietal. It differs
from the mammal in lacking an alisphenoid and a lacrimal and
in having a well developed parasphenoid and prefrontal.
Some questions remain to be solved and these are: 1. the
reality of the epiotic as a separate ossiffication and 2. the
homologies of the inner ear bones of the reptile, bird and
mammal.
LITERATURE CITED
AMADON, D. 1950 The Hawaiian honeycreepers (Aves, Drepaniidae). Bull. Amer.
Mus. Nat. Hist., 95: 151-262.
BEDDARD,F. E. 1898 The structure and classification of birds. Longmans,
Green and Co.
BITTNER, F. 1912 Uber die Schliifenregion am Schiidel der VSgel und dessen
Beziehungen zu dem der Reptilien. Archiv. f. Naturg., 78 (Abt. A ) :
1-23.
BOEM, M. 1930 Uber den Bau dcs jugendlichen Schadels von Balaeniceps rex
nebst Bemerkungen uber dessen systematische Stellung und iiber das
Gaumenskelett der VSgel. Zeits. Morph. Okol., 17 : 677-718.
BRANDT, J. FB. 1839 Beitrage zur Kenntniss der Naturgeschichte der Vogel
mit besondcrer Beziehung auf Skeletbau und vergleichende Zoologie.
M6m. Acad. Imp. Sci. St. Petersburg, Nat. Sci., ser. 6, 6: vi, 155.
BREMER, J. L. 1940 The pneumatization of the head of the common fowl.
Jour. Morph., 67: 143-157.
CUBTIS,E. L., AND R. C. MILLER 1938 The sclerotic ring in North American
birds. Auk., 55: 225-243.
CUVIER, G. 1835 Lecons d’llnatomie Compade. 2nd Ed. Crochard e t Cie.
DEBEER,G. R. 1937 Development of the vertebrate skull. Clarendon Press.
EEDMANN,K. 1940 Zur Entwicklungeschichte der Knochen im ScKadel des
Huhnes bis zum Zeitpunkt des Ausschliipfens aus dem Ei. Zeits.
Morph. Okol., 30: 315-400.
EVANS,H. E. 1948 Clearing and staining small vertebrates, in toto, for demon-
strating ossification. Turtox News, 16: 43-47.
GOODFUCH, E. 8. 1930 Studies on the structure and development of vertebrates.
Macmillan & Co.
GADOW,H., ANTI E. SELENKA 1891 Klassen und Ordnungen des Thier-Reichs, 6
(4 Abt., Anat. Th.). C. F. Winter’sche Verlagshandlung.
GRASSB,P. P. (Ed.) 1950 Trait6 de Zoologie. 15. Masson e t Cie.
GRAY, H. (ED. BY W. H. LEWIS) 1942 Anatomy of the Human Body. Lea
and Febiger.
HEILMANN, G. 1926 The origin of birds. D. Appleton & Co.
HUGGING, R. A,, 8. E. HUGGINS, I. H. IIELLWIG AND (f. DEUTSCHLANDER 1942
Ossification in the nostling House Wren. Auk, 59: 532-543.
IHLE, J. E. W. 1947 Leerboek der vergelijkende ontleedkunde van de Vcr-
tebraten. 3rd Ed., 2 vols. N. V. A. Oosthoek’s Uitgevers Mij.
JARVIK, E. 1948 On the morphology and taxonomy of the Middle Devonian
osteolepid fishes of Scotland. R. Svenska. Vet.-Akad. Hand., ser. 3,
25: 1-301.
1955 On the visceral skeleton in Eusthenopteron with a discussion
of the parasphenoid and palatoquadrate in fishes. k. Svenska Vet.-
Akad. Hand., ser. 4, 5: 1-104.
KESTEVEN,H. L. 1925 The parabasal canal and nerve foramina and canals
i n the bird skull. Jour. Roy. Soc. N. 8. Wales, Sydney, 59: 108-123.
1941 On certain debatable questions in cranio-skeletal homologies.
Proc. Linn. Soc. N. 8. Wales, 66: 293-334.
KINGSLEY,J. S. 1925 Vertebrate skeleton from the developmental standpoint.

Blakiston’s Son B; Co.
LEYMRICH,W. 1931 Der Skleralring der Vogel. Jena Zeits. Naturw., 65:
513-586.
LOWE,P. R. 1926 More notes on the quadrate as a factor in avian classification.
Ibis, scr. 12, 8: 152-188.
LUBOSCH,W. 1929 Uber den strcptognatlicn Schadel von Caprimulgus. Norph.
Jahrb., 69 (Maurer Festschrift, 2) : 96-151.
MCDOWELL,8. B., JR.,AND C. M. BOGERT1954 The systematic position of
Lanthanotus and tho affinities of the anquinornorphan lizards. Bull.
Amer. Mus. Nat. Hist., 105: 3-142.
MAGNUS,H. 1871 Untersuchungen iihcr den Bau des Knochernen Vogelkopfes.
Zeits. Wiss. Zool., 31: 1-108.
MAILLARD, J. 1948 Rccherches enibryologiqucs Bur Catharacta skua Briinn.
Rev. Suisse Zool., 55: 1--144.
MARINELLI,W. 1936 in Bolk, Goppert, Kallius and Lubosch. Handbuch der
Vergleichenden Anatomie der Wirbeltiere, 4. Urban and Schwarzen-
berg. pp. 809-838.
NEWTON,A., AND H. GADOW 1893-96 A dictionary of birds. A. & C.
Black.
NITZSCH,C. L. 1811 Ostcographische Beitrage zur Naturgeschichte der Vogel.
Carl Heinrich.
OWEN, R. 1849 On Dinornis (part 11). Trans. Zool. SOC.London, 3: 307-328.
1879 Memoir on the skull and scapular arch of the Dinornis
robustus. John vaiiVoorst.
PARKER,T. J. 1891 Observations on the anatomy and development of Apteryx.
Philos. Trans. Roy. Soc. London, Ser. B., 182: 25-134.
1892 Additional cbservations on the development of Apteryx.
Philos. Trans. Roy. SOC. London, Ser. B, 18.7: 73-84.
PARKER,W. K. 1866 On the structure of the skull in the ostrich tribe. Philos.
Trans. Roy. SOC.London, 156: 113-183.
1869 On the structure and development of the skull of the common
fowl (Gallus domesticus). Philos. Trans. Roy. SOC. London, 159 :
755-807.
1872 On the structure of thc crow’s skull. Monthly Micro. Jour.,
8: 217-226.
1873 On the development of the skull in the genus Turdus (the
thrushes). Monthly Micro. Jour., 9 : 102-107.
1874-77 On aegithognathous birds (part I). Trans. Zool. SOC.
London, 9: 289-352.
1875-79a On the morphology of the skull in the woodpeckers
.
(Picidae) and wrynecks (Yungidae) Trans. Linn. Soe. London, ser.
2 (Zool.), 1: 1-22.
1875-79b On the structure and development of the bird’s skull
(part 11). Trans. Linn. Soe. London, ser. 2 (Zool.), 1: 99-154.
1877-79 On the skull of the aegithognathous birds (part 11).
Trans. Zool. SOC. London, 10: 251-314.
PARKER,
W. K. 1880-85 On the structure and development of the skull in Croca-
dilia. Trans. Zool. Soe. London, 11: 263-310.
1890 On the morphology of the duck and the auk tribes. Cun-
ningham Memoirs, Roy. Irish. Acad., no. 6 : iv, 132.
F. R., AND T. S. WESTOLL 1941 On the evolution of the
PARRINQTON,
mammalian palate. Philos. Trans. Roy. SOC.London, Ser. B, 230:
305-355.
PETRI,CHARLES 1935 Die Skelettcntwicklung beim Meerschwein. Vierteljahrs-
schr. Naturf. Gesells. Zurich, 80: 157-240.
PYCRAFT, W.P. 1898 I n minutes of 54th meeting of the British Ornithological
Club. Bull. Brit. Ornith. Club, 7: lviii-lix.
1900-03 Some points on the morphology of the palate of the neog-
nathae. Jour. Linn. Soc. London, 28: 343-357.
RAO,M. I(. N., AND L. 8. RAUASWAMI1952 The fully formed chondrocranium
of Y a b u y a with an account of the adult osteocranium. Acta Zool.,
Stockholm, 33: 209-275.
RICHARDSON, F. 1942 Adaptive modifications for tree-trunk foraging in birds.
Univ. Calif. Publ. Zool., 46: 317-368.
ROMER,A. S. 1949 The vertebrate body. W. B. Saunders Co.
1955 The vertebrate body. 2nd Ed. W. B. Saunders Co.
Roux, G. H. 1947 The cranial dcveloprnent of certain Ethiopian L‘inseetivores’’
and its bearing on the mutual affinities of the group. Acta Zool.,
Stockholm, 28: 165-397.
SCHINZ,H. R., AND R. ZANQERL1937 Beitriige zur Osteogenese dcs Knocken-
systems beim Haushuhn, bei der Haustaube und beim Haubensteissfuss.
Denkschr. Schweie Gesells. Naturmiss., 72: vi, 161 1.+
SMITH,H. M. 1947 Classification of bone. Turtox, News, 26: 234-236.
SMITH, P. E.,W. M. COPENHAVEBAND D. D. JOHNSON 1953 Bailey’s Text-book
of Histology. Williams and Wilkins Co.
STEINEE,H. 1952 Mikro- und Makro-Evolution, der Standpunkt des Biologen.
Bericht Schweiz. Palaeont. Gesells., 45: 365-374.
STRESEMANN, -
E. 1927-34 Handbuch der Zoologie, 7 (second half Sauropsida:
Aves). Walter do Gruyter and Co.
SUSHKIN,P. P. 1899 Zur Morphologie des Vogelskelets. I. Schiidel von Tinnun-
culus. Nouv. MBm. SOC. Imp. Natur. Moscow, 16: 1-163.
1905 Zur Morphologie des Vogelskelets. Vergleichende Osteologie
der normalen Tagraubvogel (Accipitres) und der Fragen der Classi-
fication. Nouv. MBm. SOC. Imp. Natur. Moscow, 1 6 : 1-247.
TIEMEIER,0. W. 1939 A preliminary report on the 0s opticus of the bird’s eye.
Zoologica, New York, 24.- 333-338.
1950 The 0s opticus of birds. Jour. Morph., 8 6 : 25-46.
TONKOFF, W. 1900 Zur Entwicklungsgeschichte des HiihnerscEdels. Anat. Anz.,
18: 296-304.
TOEDOFF,H. B. 1954 A systematic study of the avian family Fringillidae based
on the structure of the skull. Mise. Publ. Mus. Zool. Univ. Mich., no.
81: 1-41.
WILLISTON,S. W. 1925 The osteology of the reptiles. Harvard Univ. Press.

Jollie 1957 Chick Skull

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Jollie 1957 Chick Skull

Uploaded by

Copyright:

Available Formats

T H E HEAD SKELETON O F T H E CHICKEN AND

REMARKS ON T H E ANATOMY O F THIS

A justification for another account of the bird head skele-

these is the use of whole, cleared specimens the bones of which

I want to thank Dr. Charles Peterson and the members of

fused (100 days or more after hatching). Exact ages cannot

foramina of the chicken skull are described and figured by

Fig. 1 Lateral view of 2-3 day old chick head skeleton.

the bone has allowed penetration by air sacs, particularly in

Basioccipital (figs. 3-10, 13)

Fig. 2 Dorsal view of 2-3 day old chick skull.

completely developed until its fusion with the surrounding

Esoccipital (f;gs. 1, 3-10, 12, 13, 17)

membral) early in the 12th day of incubation and gradually

Fig, 3 Veiitral view of 2-3 day old chick skull.

the posteroventral and posterior margins of the tympanic

Each exoccipital contributes a small part to the occipital

vagus and glossopharyngeal nerves as well as the blood ves-

Supraoccipital (figs. 1-10, 12)

Parker (1891 : 67) found only one in A p t e r y x , Sushkin (1899 :

dorsal margin at the line of fusion. I n the chick the supra-

Falco tinm.mcu1u.s (Sushkin, 1899, Taf. V, fig. 95), a species

Associated with the occipital ring are the bones enclosing

Prootic (Petrosal -figs. 1, 5, 6, 12, 13)

canal (enclosed in the epiotic), the posterior vertical semi-

quadrate and is separated from the squamosal by an air sac

Opisthotic (Mastoid -figs. 1, 5, 6, 9, 10, 12, 13)

The chondromembral opisthotic appears early in the 15th

semicircular canal and its ampulla and a part of the horizontal

ANT VERT SEMlClR CANA

tion of this area). During the 16th day of incubation (figs.

but later fuses with it (as in goose or Regadis-Gadoxv

bone underlying the hypophyseal fossa, so that actually the

The possibility of a “presphenoid” being involved cannot

The orbitosphenoid of the bird arises from dorsal and

Parasphenoid (figs. 1, 3-6, 17-19)

The juiictioii with the latter is detectable laterally and dorsally

Fig. 14 l'l~oto~kiicrogrnpliof x-ncvtioii throogh tlic parasplienoid rostrum show-

tixl)cwilar communis and in sonip I)ii*tls,thus contrihutes to

Fig. 15 Photomic,rograph of x-section through the cartilaginous trabccula of

the sellaparasphenoids extend down behind the eustachian

15). Each of these wings eventually forms the anterior and

Fig. 16 Photomicrograph of x-section through tlie area of fusion of the se1l:i-

part of this anterior surface devclops first and soon fuses

The basiparasphenoid centers (= tympanic bones of mam-

Fig. 1 7 Ventral view of base of craiiiuin of chick at 12th clay of iiicubntioii

Basisphenoid (figs. 1, 3, 5 , 6 , 17-19)

The basisphenoid arises from a pair of perichondral cen-

the posterior end of the rostroparasphenoid ; later these

Orbitosyhenoid (figs. 1, 2, 5, 6 , 12)

bones due t o expansion of the orbit, or brain case, or both,

8qzcninosal (figs. 1, 2, 4-6, 18, 19)

The squamosal is a membral ossification which appears late

When fully devclopetl it covei*sthe latcval aspect of the brain

I n the gallinaceous birds, not only the postorbital but also

I’nrietcrl (figs. I , 2, 4 , 5 , 7,8, 12)

Int erpci rie t m 1

at any stage, of its presence in the cranium. Staurenghy (see

Frontal (figs. 1, 2, 4,5,12,18)

Nasal (figs. 1,2,5,18)

Prefroatal (figs. 1,2,5,18)

tion of the later in relation to the lacrimal duct (see also

Nesethmoid (Perpendicular Plate of Ethmoid -