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illa Drungwara

Koomara Drungwara GoobiUa

Goobilla Belthara Koomara

Drungwara Koomara Belthara

5 This tribe inhabits the Fraser river, Derby, the Lower Fitzroy and souths
wards toward the Oakover river, Western Australia.



14


Mycoolon/ A
I B


'5


Goothanto \ g


16


Warkemon Same as


17


Koonjan/ ^
I B


18


.. (Jamagunda
J^°"e^°J»^'iGamanutU


19


Arrinda/ ^


20


Yeeda»{ J



Phratry,
A

B



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iSCfe.] MATHEWS — THE AUSTRALIAN ABORIGINES. 577

the map, comprises about two-thirds of New South Wales, the
greater part of Queensland, a wide strip through the centre of South
Australia and more than half of Western Australia.

In dealing with the tribes possessing eight intermarrying divi-
sions a table will again be employed.





Table No. 14.




JVo, on Map,


Nation,


Phratry,


Sections,


21


Wombya


Same


as in Table No. 11.


22


Ulperra «


A
B


r Ungarria, Thakamara
L Thaponungo, Jambena
r JunguUa, Thapongatee
I Joopilla, Cubadgee






L



{Boolangie, Boonongoona
, , Warkoo, Thimmemill

23 Inchalachee \

3 f KinguUa, Bungaringie

I, iNoorachee, Palyaringie

The country inhabited by the three foregoing nations is edged
yellow on the map, and comprises a large extent of the Northern
Territory of South Australia, with portions of Queensland and
Western Australia. There are several aggregates of tribes scattered
over this immense area, among each of which the names of the
sections differ more or less, but the principle of the divisions in
them all is identical. In a paper published in the Proceedings of
this Society, Vol. xxxviii, pp. 75-79, 1 particularized the eight di-
visions as found in other tribes, showing the variations in the sec-
tion names in different localties.

Nos. 24 and 25 on the map represent the Adjadurah and Nar-
rinyeri nations resj>ectively, who occupied the country fronting
Sj>encer's Gulf and on the Lower Murray river.

No. 26 is the small nation of the Kurnai in Eastern Victoria,
and No. 27 shows the territory of the Thurrawall and allied tribes
in the southeast corner of New South Wales.

In the nations Nos. 24 to 27, inclusive, the social structure is
after the tooar ox yooumree type already described by me. Adjoin-
ing the Great Australian Bight and on the west coast of Western
Australia there are also a number of tribes possessing this organi-
zation.

After much careful investigation I have succeeded in fixing on
the map the position of the blue line from A to B. All the tribes



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578 SHUFELDT — OSTEOLOGY OF THE WOODPECKEBS. [OctS,

on the west of this line practice circumcision, but in the country to
the eastward of the line the custom is not in force. The line in
yellow, from C to D, and continued on along the blue line to B,
demarks the eastern limit of splitting the urethra — a genital mutila-
tion having a very wide geographic range over the central and
western portions of the Australian continent. The other blue line,
from E to F, determines the western boundary of the tribes who
practice splitting the urethra and circumcision, neitherf of
these customs being found between that line and the coast of West-
ern Australia. From the point F, along the coastal district to
Roebourne and Condon, the rites referred to are not now insisted
upon, and it is doubtful if their adoption by the natives there was
ever universal.



ON THE OSTEOLOGY OF THE WOODPECKERS.

BY DR. R. W. SHUFELDT.
(Plate IX.)

(Read Octobers, 1900.)

Introduction.

During the past ten years I have been collecting osteological
material with the view of making a comparative study of the skele-
tons of the Ft'ci, or suborder of Woodpeckers. In time this mate-
rial became so extensive that I found I had all of the genera o^
the North American fauna represented, and had the opportunity of
examining into the osteology of many other species from different
parts of the world. Being advantageously situated with respect to
the large libraries in America, I read and investigated everything
that came to my notice upon this subject, and wrote out notes
upon the same. Occasionally I printed a brief account of some
of the osteological points of interest in this group as they came
to hand, but the main bulk of my labors in this direction have
long remained unpublished. True it is that in the Proceedings
of the Zoological Society of London I printed finally (February
3, 1 891) a short account of the osteological characters of the
Pici^ with a few brief notes upon the group, but this was
nothing more than a partial abstract based upon what had been



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1900] SHUFELDT — OSTEOLOGY OF THE WOODPECKERS. 579

accomplished up to that time. Since then all of my descrip-
tions of the skeletology of the Woodpeckers have been gathered
together, and I am inclined to believe they form the most
complete history of the skeleton in those birds, both in detail and
in a comparative way, that has thus far been furnished by anatom-
ists. For making a fair and accurate copy of my collected descrip-
tions and account of the osteology of this important group I am
wholly indebted to the kind patience and intelligence of my wife,
Alfhild, and it gives me great pleasure to thank her here for the
assistance she has rendered.

The plates and figures illustrating the present contribution have
never heretofore been published, and it is hoped that comparative
anatomists will find them useful in their work.

At least three good families of Woodpeckers are known to science
— that is, the Picida^ the Picumnidct and the lyngida — but of
these only the first-mentioned, with a variety of its genera and
numerous species, is represented in the avifauna of the United
States. Naturalists need not be reminded here by name of all
of these forms, as the majority of them are familiar to every one.
Every genus is represented by a more or less perfect skeleton in
the material I have before me at the present writing, and upon
which this account will be based ; some skeletons I have in large
series, as Melanerpes torquatus^ of which bird I collected a large
number in northwestern New Mexico in 1885. Some I have in
the young stages, beautifully exhibiting the development of the skull
and t runk- skeleton ; others, again, are more imperfect. Thanks
to the United States National Museum, I have at hand the skele-
ton of an Ivory-billed Woodpecker, but it unfortunately lacks the
skull and some few other bones. The same institution, however,
has kindly loaned me a very fair skull of Campephilus imperialis.
Numerous skeletons of various species of the genus Dry abates y of
my own preparation, are available, and I am indebted to Mr.
Samuel Parker, of Fort Klamath, Ore., for a fine skeleton oi Xeno-
picus aibolarvaius , Further, I must not forget to thank my friend
Mr. Thomas Mcllraith, of Hamilton, Ont., Canada, for a fine
skeleton of a male Picoides arcticus, I have skeletons in force of
every one of our species of Sphyrapicus^ all of my own collecting
and preparation, and by purchase I have obtained excellent skeletons
of Ceophlaus piUatus, Dr. W. S. Strode, of Bernadotte, 111., has



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580 SHUFELDT — OSTEOLOGY OF THE WOODPECKERS. [Oct 5,

kindly forwarded me skeletons of Mdanerpes erythrocephalus and
M. caroUnuSy and Herbert Brown, Esq., the well-known naturalist
of Tucson, Ariz., has, among numerous similar favors, kindly given
me the skeletons of a male and of a female Melanerpes uropygialis.
Of the genus ColapUs I have a large assortment, from quarters too
numerous to mention on this page. Mr. G. Frean Morcom, of
Chicago, and Mr. F. Stephens have presented me with alcoholic
specimens of Dryobates scalaris lucasanus^ from Lower California.

I command few or no foreign forms, but, in addition to much
other literature upon them, I have a personal copy of Prof. W. K.
Parker's memoir " On the Morphology of the Skull in the Wood-
peckers and Wrynecks " (1874).

With respect to the FicidcE, authoritative writers and ornitholo-
gists of all times seem to be of one opinion — that that family,
taken in connection with the two others mentioned above, consti-
tute a very distinct and natural group of birds with very evident
passerine affinities. The American Ornithological Union con-
siiders this group to equal an order, the Fici ; while here I treat
them as one of my suborders, which likewise is designated by the
same name.

Both Sundevall and Kessler recognized a separate group for the
Woodpeckers (Pici), while Huxley's well-known characterization
of them as his group, the CeUomorpha^ is now too well compre-
hended to render it necessary for me to reproduce it here (P. Z. 5.,
1867, p. 467). At the time that that distinguished authority pro-
mulgated the opinion to which we allude he believed that the
vomers in the skulls of some species of Woodpeckers remained dis-
tinct throughout life ; and, further, saw the nearest affines of these
birds in the Passeres (or his Coracomorpha),

In the memoir of his, to which I have just alluded in a former
paragraph, Prof. W. K. Parker wrote it as his opinion, in referring
to the Woodpeckers as a group, that *' The fact is they are like early
embryos of the Passerinae, in their palatal region arrested at a
most simple Lacertian stage, whilst in other respects they are meta-
morphosed and specialized beyond any other kind of birds. As
far as their upper face is concerned, their arrested ' maxillopala-
tines,' symmetrical 'vomers,* * septo-maxillaries ' and feebly devel-
oped turbinal scrolls entitle them to a name which shall be a
memorial of their Lacertian facial morphology. I therefore pro-
pose to call them the ^ Saurognatluz' "



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1900.] SHUFELDT — 03TE0L0GY OF THE WOODPECKERS. 581

Garrod saw the vomer of Gecinus viridis in the bone which
Parker designates as his ** medio-palatine,'* and of the "vomers"
of Parker, Garrod has said that " they look much/more like the
inner edges of the imperfectly ossified palatines." Both opinions
are candid, and simply point to a difference of opinion in a matter
of identity.*

Still another view: the distinguished ornithotomist Max Fiir-
bringer divides his order Coracomithes into three suborders, of
which the first is the Pico-Passeriformes ; and these latter are again
divided into three groups, the first of which is the Pico-Passeres,
Now he subdivides the Pico-Passeres into the Pici and PassereSy
and makes the Pici include four families, viz. , the CapitonidcB^ the
Phamphastida, the Indicatorida and the Picida? The Picida here
include the two subfamilies, the lyngina and the Picina. With
this brief recapitulation of the opinions of former writers, I will
now proceed to examine the extensive North American material
before me, describe what I find in due order, and, as usual, present
what it seems to indicate to me.

Of the Skull and Associated Bones in the PiciDiE.

As an introduction to this part of the skeleton, I select a series
of skulls taken from adult and nestling specimens of Colapies tnexi-
canus. These are very perfect and have the hyoid arches and
other bones of the sense organs associated with them.

In Colapies the premaxillary, slightly decurved throughout, is
broad at the base and gradually tapers to its apex. It is composed
of dense bone, but apparently not much more so than are the other
bones of the face. On the proximal moiety of the culmen, the
median suture is persistent at all ages of the bird, while laterally,
upon either side, the premaxillary, assisted by the nasal, surrounds
an elongated, subelliptical narial aperture of no small size. The
internarial space is largely filled in by the irregular bones of the
turbinal series and by an imperfect, though true, nasal septum.

* Compare both text and figures of Prof. Parker's memoir on the «« Morphology
of the Skull in the Woodpeckers and Wrynecks'* (Trans, Linn, Soc. Lond.y
read 1874) with Garrod's «* Note on Some of the Cranial Peculiarities of the
Woodpeckers" (Uid., 1872, pp. 357-360, and his Coll. Scientific Memoirs ^ pp.
117-119), and the figure which illustrates the latter.

' For the details of this classification, see his Untersuchungen zur MorJ>hologie
und Sy sterna tik der V'ogel, published at Amsterdam in 1888.



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582 SHUFELDT — OSTEOLOGY OF THE WOODPECKERS. [Oct. 5,

Turning to the skull of a nestling Colapies, we are to note that
the frontal processes of the bone under consideration slope gradu-
ally up on to the frontal region about equally as far as does a nasal'
on either hand. This cranio-facial region is likewise sloping in
the adult, though further on we shall see that it is quite different in-
some other Woodpeckers.

Laterally, the maxillary process of the premaxillary outerlaps the
corresponding maxillary, so that in the adult, after fusion takes
place, the two bones are indistinguishably united, and no suture
or process thereafter marks the line of union. At the roof of the
mouth, the palatine processes of the premaxillary behave in a simi-
lar manner ; the prepalatine extremity being the bone that outer-
laps the backward-projecting process of the premaxillary ut)on
either side. The apex of the premaxillary in all Picida is gen-
erally a little truncated off and squarely across.

Mesially, the premaxillary unites to some extent below willi the
nasal septum, as we so often see it in other birds wherein that par-
tition exists.

Nothing of especial import characterizes the frontal and parietal
bones in the cranium of a nestling Colaptes, Essentially they
agree with the corresponding bones in the immature specimen of
any ordinary passerine bird — a large Thrush, for example — where
the external superfices of the cranial vault is smooth and rounded..

Owing to the relatively small squamosal, however, they in
Colaptes descend far down laterally, and this feature can easily be
guessed a^ from a glance at any skull of an adult Woodpecker,
where it will be noticed that the lateral processes of the cranium,
are, comparatively near down toward the quadrate.

The frontals and parietals of an adult Colaptes do, however, pos-
sess characters not yet evident in the skull of the nestling of any.
of that genus. In the first place, there is the denting of nearly the
entire external surface of these parial bones, which is caused b/
the apices of the quill-butts of the capital feathers. Again, mre
note in this region, in the, adult, the double impressed groove,
leading from the supraoccipital prominence by a gentle curve to
the posterior periphery of the right narial aperture. This double
gutter lodges, as we know, the free ends of the hyoidean apparatus
during the life of the individual. Adult Colaptes also exhibit a
slight tilting up of the postero-superior orbital peripheries, the
margins being sharpened and a few perforating foramina occurring



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190a] SHUFELDT— OSTEOLOGY OF THE WOODPECKERS. 58S

just within their borders, one behind the other down the line.
The frontal region between the rims of the orbits above is but
moderately broad, and barely concaved.

A nasal bone in the skull of the young Colaptes has the usual
form seen among the higher passerine types ; it is of the holorhinal
pattern, and makes the usual articulations with the surrounding
bones. As the bird matures a diminutive process is seen to appear
upon its posterior margin between the anterior apex of the frontal
and an abutting process thrown up on the part of the corresponding
maxillo-palatine. Anteriorly it sends a jagged horn of some size
forward into the anterior rhinal space. A true lacrymal does not
develop in Colaptes, The pars plana is large, very complete and
of a quadrilateral outline. Postero- externally it sends backward
from its angle a long, pointed os uncinatum that normally bears
upon the inner side of the jugal bar. Colaptes^ agreeing with all
our United States Picidcty exhibits considerable ossification of the
turbinal series of bones, although there is a marked simplicity of the
arrangement of them. One turbinal seems to be free upon either
side, both in this and in other species. Parker has paid consider-
able attention to the turbinals of the Pici^ and in briefly alluding
to what he saw in other species, in the ninth edition of the Ency-
clopadia Britannica (article "Birds,** p. 717), he says that "the
' inferior turbinal,' which has three coils in Rhea and Tinamus and
two in most birds, is in Gecinus merely bi-alate ; in Junx it makes
less than a single turn, whilst the alinasal turbinal of that bird has
two turns and that of Gecinus one. Gecinus is in all respects the
most specialized, Picumnus the most embryonic and Junx the
most passerine of the Celeomorphce,'*

In adult specimens of Colaptes the interorbital septum is usually
entire, though it often shows a very minute central vacuity. When
we come to examine other species of Woodpeckers, later on, we
will see that this foramen in them is larger and more constant.

The infraorbital bar or zygoma is made up entirely of the juga)
and maxillary, the quadrato-jugal not being present. It is straight
and slender, being abruptly enlarged at the posterior o: jugal ex-
tremity, where it is hooked inward to articulate with the quadrate.

Nestling Colaptes show the development of a fine pair of orbito-
sphenoids, occupying their usual sites at the back of the orbits;
and within these cavities, above either orbito-sphenoid, we observe
the free edge of the frontal showing its advancing ossification. At



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o8db SHUFELDT— OSTEOLOGY OF THE WOODPECKERS. [Oct. 5,
certain points this is peculiar, as little islets of bone occur along



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rt .• rt S



7



♦* s ••*

-s > i 3 S
11° II

ju »^ fe 2 -^

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« g ■•^ e

.3i o « g *-.
> • cu-^ »^
►• ;t « 3

C ^ • >

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<3 X « * «
^ ft' .S k S

^ 8-1 !^ ''*
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S ^^ ITS

- r£ S. :5' • -



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1900.] SHUFELDT — OSTEOLOGY OF THE WOODPECKERS. 585

this free edge, which are afterward absorbed by the main bone.
This process occurs in other parts of the skulls of Woodpeckers.

Passing to the base of the skull, we see a number of points of
interest in ColapUs, and some few features that are not present in
the general run of ordinary existing birds. The basitemporal re-
gion is smooth, broad and deep antero-posteriorly. Nothing
especially noteworthy characterizes the large foramen magnum,
with its relatively small condyle, nor the foramina in front of it
upon either hand. A supraoccipital prominence is also present,
being but fairly developed, and never pierced by foramina in the
adult individual.

Either tympanic bulla is cowrie shaped, showing a characteristic
antero-posterior slit, and, according to Parker, this unique feature
oi the Woodpeckers is brought about by one of these bullae being
formed by not only the corresponding exoccipital, but by two, or
more often several, tympanices (usually three). The basitemporal
also enters into its formation (see Fig. i).

A quadrate is of good size comparatively, with its orbital process
stout and truncated at its apex. The "mastoidal limb" is short,
and the mandibular facets peculiar. Either pterygoid is noted for
its pointed and lengthened meso-pterygoid, which, when in siiUy
reaches forward to the palatine spur of the same side. On the
upper edge of a pterygoid a prominent and pointed muscular pro-
cess is developed ; it has the same direction forward and inward as
has the shaft of the bone. This shaft is thin and much compressed,
while the quadratal head is small and inconspicuous. When artic-
ulated, the palatal heads do not meet mesially, or they may just
barely touch each other. Occasionally we find in old specimens
-of ColapUs minute osseous " prickles " at the usual sites of the
basipterygoidal processes, and they represent aborted basiptery-
goids. They are more prominent in certain other Woodpeckers of
our avifauna, but it is only rarely that the corresponding process is
ever seen upon the pterygoid. It is present, though very rudimen-
tary, in a specimen of the Pileated Woodpecker before me.

The inferior border of the rostrum is rounded and not very thick
transversely. It is very sharp in front, and below is usually carried
-out as a little spine. Either palatine is narrow, especially ante-
riorly beyond the maxillo-palatine, or what is known as its pre-
palatine portion, Distally this extremity is underlapped by the
palatine process (of the same side) of the premaxillary. Poste-



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586 SnUFELDT — OSTEOLOGY OF THE WOODPECKERS. [Oct*,

riorly the posteroexternal angle is obliquely truncated, and the
head for the corresponding pterygoid is completely aborted, the
palatine being simply drawn out at this end. About midway up its
mesial border it develops a conspicuous interpalatine process,
directed forward, while another free process, also directed forward,
springs from that point on the bone where the bifurcation of the
vomer articulates in a great number of ordinary birds. This last
process, so variable and so conspicuous in nearly all Pici^ I here
designate as the palatine spur. During ossification of the mesial
and external margins of either prepalatine minute islets of bone are
left, and these may persist or they may not persist during the life
of the individual.

The vomer, mesial in position, is situated rather far back be-
tween the palatines! and is non-bifurcated posteriorly. It is pointed
in front and pointed behind, and rests for nearly its entire length
upon the nether aspect of the sphenoidal rostrum. In some Wood-
peckers it is quite compressed in the vertical direction and of
somewhat of a lozenge-shaped outline. That it is not forked pos-
teriorly need not surprise us, for that also is the case with the semi-
rudimentary vomer in Geococcyx and some other birds.

The lower margin of the mid-lying nasal septum is long and
scraggly ; while above, the laminated portion of the bone is not
completed up to the premaxilline roof. A maxillo-palatine is of
considerable size, broad, horizontally flattened, perforated upon its
nether side by foramina, and finally it sends backward from its
mesio-posterior angle a blunt-like process. This last feature is
absent in many Pici^ and here in Coiapies the inner margin of a
maxillo-palatine barely comes in so far as the outer border of the
corresponding palatine, and so these bones are widely separated
from each other in the middle line.

In Colaptes the mandible is of the typical V-shaped pattern, long
and narrow, with a very shallow symphysis. Its ramal sides are
rather shallow, upright and with rounded superior and inferior
borders. A ramal vacuity is absent, or reduced to a mere pinhole
or smaller. The angular extremities are rounded, and offer no
special process on either side. This jawbone is pneumatic, and its
hinder articular ends present the usual ornithic characters, with
perhaps an arrangement of the mandibular facets peculiar to the
Pici, but departing but little from what we find in almost all ordi-
nary avian types. In the adult all the sutural traces among the
splint-bones composing the mandible have disappeared.



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two.] SHUFELDT — OSTEOLOGY OF THE WOODPECKERS. 587

A very curious condition of the sclerotal plates of the eyeball
exists in Colaptes and all other American Pici wherein I have exam-
ined them. These plates very thoroughly fuse together, although
in the nestling at the time it leaves the nest they are still separate.
In the case of the sclerotals of a specimen of the big Imperial
Woodpecker before me this fusion is so perfect that all the sutural
traces among the plates have well-nigh disappeared entirely. Such
a condition is not common among highly organized birds.



Fig. 2. Superior view, natural size, of the skull of the Imperial Woodpecker
(C imperialist Drawn by the author from specimen No. 1464 of the oste-
ological collections of the United States National Museum.

The present writer has not paid any very special attention to the
bones of the ear in the Fici, but Parker has said of them for Ge-
cinus viridis that "The stapedial apex of the largely aborted
second postoral arch has some peculiarities of importance. The
true stapedial or periotic portion is rather large and roughly oval,
the side towafd the * opisthotic * bar separating the fenestra oval is



Online LibraryHague (Netherlands). Conférence Internationale surProceedings / Linnean Society of New South Wales, Sydney, Volume 18 → online text (page 54 of 68)