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IFrom
The
'
Tk.u»-sactions of the Zoological Society 0/
London
',
— Vol. xv. Part
December 1900.]
v.,
A
Monograph of the_Genus Casuarius./ £1/ the Hon. Walter Rothschild, Ph.D.,
F.Z.S. With a Dissertation on the Morphology and Phyhgeny of the PAL^OGNATHiE
(Ratitj; and Cryi'Turi) and Neognath^ (Carinat^).
By W. P. Pycraft.
VI.
Kcceived nnd read Juno 20, 1899.
[Plates
Part
By
I.
—A
Monograph op the Genus
CASUARIUS
Genus
1790. Caxuarius Brisson, Orn.
Handis
u.
Mem.
de
I'lnst.
519.
New
Guinea with
the islands of the
Aru group,
and inhabits the Papuan subregion,
New
Britain,
Northern Queensland, and the island of Ceram
many
evidently separable into a great
but owing to their large
living specimens that are
p.
Hilfsbuch, pp. xxxxiii, 452.
the islands in Geelvink Bay, Salwatty,
;
iii.
the typical genus of the group Casuarii of the PalceognathfV
(Batitce of former authors),
species
Biiss.
v. p. 10.
1801. Rhea (iion Latham, 1790) Lacepede,
The genus Casuarius
Casuarius.
Hon. Walter Rothschild, Ph.D., F.Z.S.
the
1842. Hippalectryo Gloger,
XXII.-XLV.]
to
The
Moluccas.
local forms, apart
size, to
brought
in the
i. e..
Cassowaries are
from the few very
distinct
good
the uncertainty about the localities of the
Europe, and to the disappearance after death of
the most characteristic coloration and structure of the bare skin on the head and neck,
(jur
knowledge of the species
been able
is
to study is very large
doubtless
The
compared with that available
several years I have tried to procure a great
as well, and I
limited.
still
have been able to
number
place numerous
Mr. Pycraft, whose a])pended paper
will, I trust,
material which
in
I
have
most Museums.
For
of living specimens and of skins
fresh bodies
and skeletons before
be of great value for the study of the
anatomy of the Palwognatha?.
Outwardly the
peculiarities
The
genus
Casuarius
may
easily
be
recognized
by
following
the
:—
bill is short,
generally shorter than the head, laterally compressed, strong, and
with the culmen curved downwards near the
middle of the maxilla,
in
tip.
The
nostrils are
placed in the
a wide membranous groove, and open anteriorly.
Head and
neck bare and mostly brightly coloured, tlie skin much carunculated and wattled in
The forehead is ornamented with a more or less developed helmet of
various places.
various shapes.
Wings
any indication of webs
VOL. XV.
— PART
v.
quite rudimentary, with about five round black shafts, without
in the place of remiges
No.
1.
December, 1900.
;
no rectrices apparent.
The metatarsus
s
HON. W. EOTHSCHILD ON THE GENUS CASUARIUS.
110
is
shorter than in most other Ratitse, very robust, and covered in front with hexagonal
and transverse ones near the
scutes,
curved, and short claws
is
Avhich
the after-shaft
is
covered with
when
nestling,
hatched,
The
shaft.
is
hair-like featiiers, in
stiff,
old birds are black, the
striped longitudinally above.
eggs are about six to eight in niimber, Avith a strong and very coarsely granulated
When
surface.
fresh they are evidently all of a light green colour, but
they become
to the light
first
The male alone
coloured.
Herr Schalow (Journ.
f.
ju'iucipally
more
cream-
incubates.
;
but his key
based on the colour, which
which
when exposed
bluish, then greyish, and at last almost
Orn. 1894) has attempted to
eggs of the various species
shell,
The body
as long as the principal
is
young ones brown, the
The
the inner toe with a long, straight, powerful, pointed claw,
;
a dangerous weapon.
Avhich
Toes three, the two outer ones with obtuse,
toes.
the
is
a total
is
same
make
failure
in all,
key to distinguish the
a
and misleading, being
and on the transparency of the
is not specifically different.
All Cassowaries are inhabitants of forests, while the rest of the large living Palceognat]ice{ox Ratitce) are denizens of steppes
all sorts
of vegetable matter and fruits
small birds
when
they
come
sleeping from sunset
The
till
but they also pick up insects and any creeping
;
They
across them.
swallow quantities of stones and gravel
Their food seems to consist of
deserts.
In captivity, at
in their way.
thing that comes
and
least, tliey kill
and devour chicks and
also, like Ostriches,
They
to assist digestion.
Kheas, and others,
are entirely diurnal,
morning.
voice of the Cassowaries
is
a curious sort of snorting, grunting, and
bellowing,
usually not very loud, and differing according to the species.
Their temper
is
generally sullen and treacherous, and they are extremely pugnacious,
even the different sexes often fighting at other seasons
Exceptions are rather rare, but there
which
is
quite tame, and
to follow the
was always
an adult Cassowary, now in the Society's Gardens,
so, since I
keeper who fed him like
become somewhat
It
is
than the breeding-season.
a
had him
dog when
I
;
while another young bird used
had him
in Tring,
but has since
shy.
seems that the Dutch navigators
in
1596 were the
Europeans who became
first
acquainted with the Cassowary, at least there does not appear to be any indication
that the Portuguese,
the bird.
who
visited the East-Indian Archipelago long before,
came
For more than two and a half centuries there was no certainty
the existence of more than one species, but in 1854 Mr.
across
as
Thomas Wall procured
to
a
specimen of Casunrius castiarius australis near Cape York, which was described by his
brother, for the
first
(made from memory)
time, in
is
as
the
bad as
it
'
Illustrated
ran be.
Cassowary, in 1800 Blyth and Sclater
Sydney Herald
In 1857 Gould
described
best general accounts are those by
Salvadori in his
;
but the description
described the
two new
the described forms have constantly been increasing,
'
species,
New
Britain
and since then
now number liO. The
monograph of the genus in the
till
they
—
—
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
'
Memorie
Accademia
della
di Torino,'
aud
famous
in his
'
Oniitologia della Papuasia,'
pp. 473-508.
iii.
The Cassowaries
are easily divided into three
j^n'oups,
HI
as follows:
group, containing two species, Casuarius bicarmiGulatus and
—
The
^1.
C. casiiarius
with
typical
its
seven
The one-wattled Cassowaries, containing two species, C. philipi and
C.
nappendicidatns, with ionr subspecies
The Mooruks, which number four
o.
species
C. papuanus with two subspecies, C. picticolUs with two subspecies, C. bennetti
subspecies
2.
;
11
;
with two subspecies, and
and anatomically.
may be roughly designated
thus
:
....
two wattles on fore-neck
laterally,
Casque depressed posteriorly, sometimes compressed
Typical Group.
one
laterally,
One-wattled Cassowaries.
wattle on fore-neck
Casque depressed
In
the
species
fact
group the species
Casuarius casuarius and
and slope of the casque
in the height
subspecies
;
exist
these
in
casuarius
Also we often find that males of one given subspecies
there
first,
because there
enormous casques, while
casque at
have
no doubt that certain differences
is
not be placed on these
considerable individual variation; secondly, because
and, thirdly, because the period of growth seems variable
;
captivity).
in
least
is
much weight cannot and must
but too
;
sexual variation
is
There
two characters among the subspecies and species of the Casunrhis
section
differences:
have seen young birds
I
fully adult
birds with
in
still
brown plumage with
hardly any
black plumage showed
all.
to
thank
Finsch,
Drs.
Gestro,
and Heck,
Giglioli,
Count
Salvadori,
me
specimens,
Dr. Sclater, Mr. Blaauw, and others, for their kind help in lending
drawings, &c.
Key
,'
-I
'
Plumage
striped.
to the Species
Immature.
Pkimage
Adult
black.
and Subspecies of
Genus Casuarius.
the
C/iicks.
Plumage
'& brown.
/Two
wattles on fore-neck
2.
;
casque not depressed posteriorly,
but compressed laterally
•'^
j
2
•(
One
wattle on fore-neclc
;
casque depressed posteriorly or
S.
compressed laterally
I
I,
vary
to the
much higher and more erect casques, while the females have the casque
lower and sloping back over the occiput.
1
""
remains that we often have greater differences among individuals of one subspecies
show, as a rule,
I
its
and although in the " Key
have stated that certain forms are distinguished by a high or low casque, the
I
than between two subspecies.
(at
The Mooruks.
no wattle on fore-neck
posteriorly,
typical
enormously
The
Mr. Pycraft's memoir will detail the anatomical features.
external characters
Casque compressed
These three groups are readily separable externally
C. /or/re.
No
wattle on forc-ueck, casque depressed posteriorly
.
.
11.
S 2
HON. W. EOTHSCHILD ON THE GENUS CASUAKIUS.
112
C. blcarunculatus.
Wattles wide apart on sides of neck
3
Wattles close together on front of neck
/
Size large
;
Size large
;
4.
wattles very large, above 5 inches long
wattles
medium, 3 inches or
less,
or small
5.
.
.
.
.
6.
.
I
4
Size small
•{
casque low, sloping, and less erect
;
sides of
;
neck anteriorly bluish purple, posteriorly red
.
.
.
C. casuarhis.
.
C. casuarius sclateri.
I
Size small
\
;
casque very high
7.
Sides of neck anteriorly bright blue, posteriorly scarlet
||
I
.
\
Sides of neck anteriorly bluish purple, posteriorly scarlet
(
Casque low but very
!
of neck purplish
;
sides
(' casuarius violicolUs.
mauve
C. casuarius intensus.
Sides of neck entirely blue
anteriorly blue, posteriorly scarlet
\
Sides of neck
\
Fore-neck blue, wattle crimson and blue
;
....
Fore-neck
!
iu
blue
front,
orange-scarlet
at
the
.
.
...
(
.
.
Casque compressed laterally, fore- and hind-neck and sides
of the neck uniform orange
Casque low, the blue not extending below chin, naked sides
\
i
(
'^
uuappeHdiculatus rufutinctus.
9.
10.
C.
umppendiculatus aurantiacus.
C. unappendiculutus.
of neck yellow veined with red
I
C. philipi.
C-''
Fore-neck orange or yellow, wattle yellow or livid
Casque depressed posteriorly, not compressed laterally
I
casuarius salcudorii.
sides
wattle blue
I
C
casque very
high, compressed laterally, depressed posteriorly
I
9
C. casuarius beccarii.
f
'
w
sides of
Casque low, not very large, wattles separate at base
I
^
;
neck anteriorly blue, posteriorly scarlet
1
8
large, wattles joined at base
C. casuarius australis.
.
Casque very high, the blue extending on to the throat, naked
1
sides of
1
neck crimson
•
(J-
unappendiculat us occipitalis.
1~-
Fore-neck blue
j
^^
[
^^
C. loricB.
Hind-neck
13.
scarlet
J
I
14-
Hind-neck blue
side of throat below ear surrounded
Lar^e black patch on
by a broad pink band
1
13
Fore-neck red, throat pink
I
[
f
black and pink patch on sides of throat
Head and neck blue, sides of neck black with pink between
Xo
the wrinkles
C.
pupuanus edward^i.
C.
papuanus.
t,'-
bennetti.
j
1
^^j
Head and neck
of
blue,
round patch on fore-neck and sides
C. bennetti muvuhitus.
neck rosy purple
Head whitish
blue, fore-
and hind-neck blue, sides of
I
1
j
^^
j
neck purplish red
15.
Fore-neck entirely blue
C. piclicollis /itcki.
Fore-neck with red patch iu centre
C. picticollis.
II
;
.
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
1.
1603.
Casuarius CASUARIUS
Jrw
Erne Aldrovand., Oru.
obtained one in Java
1605.
1635.
Common
(L.).
iii.
541
p.
Coram Cassowary.
or
113
XXII.)
(Plate
(Banda, brought to Hollaud by Scheliinger, who
very bad figure)
:
Emeu Clusius, Exotic, lib. v. p. 98 (good figure,
Emeu Nieremberg, Hist. Nat. p. 218 (fig.).
1666. Casuar Olearius, Gottorffische
taken from sanae specimen).
Kunstkammer,
23,
p.
pi.
xiii.
Gottorff
(alive in
;
figure
copied from Clusius).
&
1666-1699. Casoar Perr.
{teste Salvadori)
Emeu Ray &
Ac. Sc. depuis 166ei-1699,
iii.
2, p.
155, pis. 56, 57
;
1770. Buflbn, Hist. Nat. Ois.
1676.
Mem.
Duvern.,
464.
p.
i.
Willughby, Orn.
p. 105, pi. xxi.
Oud eu nieuw Oost-Indien, iii. (Amboina), p. 298.
& Derham, Nat. Hist. B. ii. p. 56, pi. Ix. (alive at George
1726. Kazuwuris (Ceram), Valentyn,
1738. Cassowary or
Emeu
Albin
Tavern,
Charing Cross).
1745.
Emeu
Barrere, Orn. spec. uov. class. 3, gen. xxxviii.
1750. Caseurius Klein, Hist. Av. Prodr.
Avium Genera, no.
Emeu Worm, Mus. Worm. p. 292.
1752. Cela Moehring,
1755.
1756. Avis
Emeu
56.
Johnston, Theatr. Uuiv. de Avibus,
1758. Strut/do casuarius Linnteus, Syst. Nat. ed.
1766.
Id., Syst.
Nat. ed.
xii.
1788. Gmelin, Syst. Nat.
p.
i.
1760. Casoar Brisson, Oru.
i.
265
p.
10, pi.
vii. p.
1828. Lesson, Voy. Coq., Zool.
i.
p. 7, pi.
Hauptai't.
p.
ii.
664
(partim
fig. 1
p. 2, pi.
345,
1819. Merrem, Abhandl. Berl. Akad.
v, p.
p.
V.
in
Ersch
&
f.
Orn.
225
2, p.
pi.
432,
1861. Rosenberg, Journ.
f.
Orn.
p.
1862. Crisp, Proc. Zool. Soc. Lond.
(egg)
;
B. 11.
2;
fig. 1
pi.
29
;
348;
p.
;
1860. Sclater, Proc. Zool. Soc. Lond. pp. 210, 250
1860. Des Murs, Tr. d'Oologie, p. 362
1
;
365
p.
;
fig.
p. 4, pi. iv. fig.
Gruber's Encycl. xv.
Homeyer, Journ.
;
179;
1819. Stephens in Shaw's Gen. Zool. xiv.
p. 77, pi.
!)
p. 9.
1816. Vieillot, Nouv. Diet. Hist. Nat.
Merrem
{gO(jtl fig.
;
!)
i.
i.
1859. A.
;
105, 105 a
1, pi.
711 (partim
pt. 2, p.
ii.
1790. Casuarius galeatus Bonn. Euc. Meth.
1826.
155
199;
1873. Schlegel, Mus. P.-B., Struthiones,
ii.
p.
56.
fig. 2.
i.
ix.
1845. Thienemaun, Fortpfl. ges. Vogel,
1825. Vieillot, Gal. Ois.
i.
pi.
726, no. 2.
v. p.
1790. Casuarius emeu Latham, Ind. Orn.
1831. Id., Traite d'Orn.
x.
ISO,
p.
;
1763. Casuarius Frisch, Vorstell. Vog,
1817. Dum., Diet. Sc. Nat.
1.
p. 17.
;
;
45;
p.
137;
1862. Sclater, Trans. Zool. Soc. Lond.
iv. p.
358,
fig. a, p.
360
(note), pi. 71
;
bird
and egg).
HON. W. KOTHSCHILD ON THE GENUS CASUARIUS.
114
1862. Schlegel, Jaarb. zool. Genootsch. Nat. Art.
1863. Sclater, Proc. Zool. Soc. Loud. p. 234
Mag.
p.
196
;
1864. Schlegel, Diereut., Vogels, p. 238;
1866. Sclater, Proc. Zool. Soc. Lond.
1867. Id., op.
cit. p.
1869. Wallace,
1
168;
p.
79 (hatcbed in Society's Gardens)
Malay
Arcliip.
156
p.
ii.
;
;
1871. Flower, Proc. Zool. Soc. Loud. pp. 32-35;
1873. Garrod, Proc. Zool. Soc. Lond.
1873. Rosenberg, Journ.
f.
Orn.
614;
p.
390
p.
;
1875. Sclater, Proc. Zool. Soc. Lond. pp. 86, 87;
1877. Harting, Ostriches
&
Ostr. Farm. p. 103, plate;
1879. Rosenberg, Malay. Arch. p. 323;
1881. Sclater, Voy. 'Challenger,' Birds,
1881. Salvador!,
Mem.
1882. Id., Orn. Pap. e Molucc.
iii.
1891. Thebault, Bull. Soc. Phil.
1891. Salvador!,
Mem.
1894. Schalow, Journ.
64;
p.
R. Ac. Sc. Torino, (2) xxxiv.
p.
(8)
479
iii.
R. Ac. Sc. Torino,
f.
Orn.
18 (egg)
p.
2
;
;
p.
592
;
1811. Casuarius casuarius liliger, Prodr.
Mamm.
et
263 (one menagerie bird only
Av.
1842. Hippalectryo indicus Gloger, Hand- u. Hilfsb.
p.
1872. Hippalectryo caauarius Suudev., Tentamen,
p.
in
.Museum).
p. 247.
452;
1844. Casuarivs orienfaUs S. Miill., Land- en Volkenk.
p.
109 (partira
!).
152.
Gulliv., Proc. Zool. Soc.
!)
fig.
213;
(2) xlii. p.
(3) viii. p.
(sic
i.
198-210;
1896. Oustalet, Nouv. Arch. Mus. Paris,
1875. Casuariiis javanensis
pi.
;
pp.
1895. Salvador!, Cat. B. Brit. Mus. xxvii.
188,
p.
Lond.
jjp.
478, 488.
1879. Casuarius beccarii (non Sclater!) Pelzeln, Ibis, 1879, pp. 376, 377
;
1881. Id., Ibis, p. 401 (egg).
Native name
Ceram,
or,
:
what
is
more
Adult. Bill long
compressed
Suwuri by the Malays
called
word used
likely, a
but
;
is
it
not certain
this
for all Cassowaries in the trade
is
name on
a local
and of Papuan
origin.
casque large, not very high, and soinewliat sloping backwards,
;
laterally, of a
dark brownish horn-colour.
becoming darker on the upper part of the hind-neck.
Chin, throat, and fore-neck dark blue.
scarlet.
if
On
Head and
occiput Nile-blue,
Lower two-thirds
of hind-neck
the lower fore-neck are two large
lappet-shaped wattles, joined at their base, rugose, granulated, and of a deep pink
Naked lower
colour.
])osteriorly.
Lores black
sides
;
iris
of neck
bluish
dark brown
;
legs
purple anteriorly and
brownish grey.
bright
Plumage
scarlet
black.
Size
medium.
Total length about 1| m.,
inner toe 70-78
Young
brown.
sides of
bill
from gape 98 mm., tarsus 250
to
275 mm., claw of
mm.
(two-fifths grown).
Head and
Plumage of various shades
of yellowish
brown
to rufous
fore-neck dull leaden blue, hind-neck dull yellow, naked lower
neck a mixture of dull blue and
livid purple.
HON. W. ROTHSCHILD ON THE GENUS CASUAKIUS.
115
J o?/wy (tliree-fif'ths grown), riumage yellowish or rufous brown mixed with black.
Fore-neck dull indigo-blue; head and occiput pale dull blue; hind-neck dull orangered, naked lower sides ot neck blue, ])osteriorly mixed livid purple and dull red.
Young
(full-grown).
Plumage
mixed with a few rufous
black,
Fore-neck
featliers.
iiubgo-blue; head and occiput ))ale l)lue
hind-neck scarlet, naked lower sides of neck
blue anteriorly, passing through plum-purple to scarlet posteriorly.
;
Head and neck
Chick.
rufous,
paler
below
three
;
broad
stripes
back, one
on
irregular stripe on each side reaching from the anus to the wing, and two other lateral
stripes
extending to the thighs, the
last
two broken up into irregular blotches.
Hah. Geram.
As shown
genus.
in the
number
In the year 1595 a
and sent out four
bring
synonymy, the Ceram Cassowary
home
vessels to
of merchants in
open communication with the Eastern Archipelago and to
In December 1596 the ships were
spices and other valuable merchandise.
anchored at tSydayo,
of the ship
in Java,
and
it
was there that Jan Jacobsz Schellinger, the skipper
Amsterdam,' was i)resented with
'
Java from Banda Island.
the chief of Sydayo.
wonderful " large fowl
known member of the
Amsterdam formed a company,
the oldest
is
which had been brought
a Cassowary,
This was a day or two before Schellinger was murdered
Although the ship 'Amsterdam' was
" \\as
landed alive at Amsterdam
and
left
burnt,
to
l)y
the
brought on board of one of the other vessels and was
1597.
in
It
was
at first exhibited
to the public for
some
months, then came into the hands of Count George Everard .Solms, of s'Gravenhage,
who kept
for a long time at
it
the Hague, and afterwards presented
Prince Ernestus of Cologne, who, again, subsequently gave
Count Solms, before parting with the
the Second.
picture
made
one of
its
of
it,
feathers,
A
other works.
from which the very good figure
and an egg, was taken.
figure
despicable work
'
description
have been the
had an excellent coloured
in Clusius, representing the bird,
it
was
first
1666 Olearius,
In
in the diary of
observed by Europeans, .ind
in
his
somewhat
Die (jottorfhsche Kunstkammer,' assures us that one had been kept
alive at Gottorff in Schleswig
is
Emperor Rudolphus
wretched figure of the same bird was also published
reproduced by Aldrovandus.
is
to the
the Elector,
to
This figure has been copied into several
the long and dangerous voyage during which
this
it
bird,
it
;
but he does not say how
it
was procured.
very amusing, but short and not very accurate.
first to
give the actual liome of this bird, which
some Dutchmen found
it
sitting
Olearius's
Valentyn (1726) seems
is
Ceram.
He
to
says that
on three eggs on that i^land as long ago as 1660.
Valentyn also says that the Cassowary occurring
in
Aru
differs
from that of Ceram.
Other writers have stated Sumatra, Ceylon, and all sorts of places to be inhabited by
" Habitat in Asm, Sumatra, Molucca, Banda."
Cassowaries, and even Linnaeus said
:
Up
to the present day erroneous statements respecting the
have often been made, but
inhabited by a (Cassowary.
it is
evident that of
all
home
of Casaariuts casiiurin.t
the Moluccan islands only
Ceram
is
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
116
Linnaeus,
1758, bestowed
in
our present
Struthio casuarius, and diagnosing
He
busque nudis.'"
then
;
adds
finally
coUum
a tergo
:
this
calling
bird,
it
tridactylis, vertice paleari-
Dodart, Bay, Clusius,
quotes Olearius, Albin,
Willughby, Aldrovandus, and
Palearia duo sub collo
it
name on
specific
as " Struthio pedihus
Bontius,
" Callus elevatus quasi diadema verticis.
Ungue intermedia majore
nudum.
;
ova
in
the
ferit
puuctis excavatis."
The
species
interior of
admitted by Wallace and Rosenberg to be not
is
Ceram, but
approach, and
it
less trustworthy,
is
it
extremely shy and
seems that no European naturalist has ever seen
we know about
Ail
that
travellers say
all
but
it
seems certain that
eggs are said to be three to
A
three eggs only.
we cannot
believe that
its
pair,
at
Tring Park,
The
to
six
clutch can regularly contain
young Casuarius casuarius was hatched
in 1867, but observations respecting the time of incubation
were apparently not made.
in its wild state.
it
feeds principally on fallen fruit.
it
number; but we have had,
five in
eight laid by one female, so that
to
difficult
comes from native sources, and may be more or
life-history
its
at all rare
the Society's Gardens
in
and other interesting points
have never succeeded in getting a male that would
I
although I have had a female which laid many eggs.
The egg
by
described
Schalow (Journ. f. Orn. 1894, p. 19) is faded. There is, I am convinced, not a single
form of Cassowary that lays " pale greyish-green " eggs with " dark grey-green" glazed
In any case Casuarius casuarius lays light apple-green eggs, with darker
granulations.
They
bright green corrugations.
are elliptical, the thickest zone about the centre,
though very often a thick and a thin end can be distinguished.
glazed granulations are
all
more
The measurements are:— 150
145
:
85
:
The
elevated
or less connected, forming a labyrinthic network.
142
94,
93, 140
:
90,
:
131
:
90,
129
:
87,
123
:
84,
mm.
The eggs
Owing to their size it is difficult to boil them well in
number of persons to finish one but for scrambled
and other purposes they are, like those of the Rhea and Emu,
are excellent food.
the shell, and besides
eggs, omelettes, cakes,
it
requires a
;
most excellent.
2.
Casuarius casuarius beccarii
Beccari's Cassowary.
Scl.
1875. Casuarius beccarii Scl., Proc. Zool. Soc. Lond.
Islands
!)
p.
87,
1878. A. B. Meyer, Journ.
Mem.
f.
Orn.
p.
300
R. Ac. Sc. Torino,
IS82. Id., Orn. Papuas. e Molucc.
1894. Schalow, Journ.
f.
Orn.
p.
iii.
p.
;
Salvad.
(?)
2 oa
p.
86 (Vokau, Ani
197
(2) xxxiv. p.
484 (partim
21 (egg, Aru)
Casuarius bicarunculatus (non Scl.
!)
;
Ann. Mus.
:
Civ. Genoa,
(partira
:
xii. p.
Aru
only)
Civ.
Gen.
421
Aru)
;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 596 (partim
1875.
1,
XXIII.)
;
1877. Forbes, Proc. Zool. Soc. Lond. pp. 307, 316 (anatomy)
1881. Salvad.,
figs.
(Plate
:
Beccari,
Aru).
Ann. Mus.
vii. p.
717 (Aru).
—
—
HON.
1881.
ROTHSCHILD ON THE GENUS CaSUAEIUS.
^Y.
Casuurii>s galeatus
(?j
bought
in Ternate
(iioii
—
B. Brit. Mus. xxvii.
1881.
Bonn.
Zool. Voy.
Scl.,
!)
seem to be young of
C.
c.
ge.s.
Orn.
Aru
an egg without history
Advlf. Plumage black
from
tlic
64
p.
Aru Islands
(youiii;,
cf. Ciit.
;
p. ."jD?).
C'asuarius sp. A. B. Meyer, Zeitschr.
al)out
Challenger,' Birds,
heccarti
liowever, several Cassowaries inhabiting the
(?)
'
117
i.
p. 2[)G
(egg from
Islands, there
is
Aru— as
no certainty whatever
!).
casque very high, broad and massive, dark brown
;
there mx-,
bill
;
long
Head and
and not pointed.
occiput pale greenish blue.
Upper part of hind-neck
deeper blue, followed by a space of mixed blue and red, ])roducing a somewhat livid
Lower part of hind-neck
pule purple eflect.
dark blue.
Chin, throat, and fore-neck
scarlet.
Wattles large, broad, and very conspicuously joined
at
base, of a
deep
up in a narrow line nearly to below the ear.
of neck mixed red and blue.
A very large bird, but 1 have not
fleshy-red colour, wliich colour runs
Naked lower
been able
Young
sides
measure a
to
dead example.
fully adult
(three-fourths grown). Fore-neck indigo-blue, head and occiput pale blue
;
hind-neck orange-yellow, with a purple patch between blue and orange. Naked lower
sides of neck, anterior half dull blue, posterior lialf orange-yellow.
Plumage brovpn.
Young (about
full-grown).
Same
only colours less bright and plumage
as adult,
brown.
Hab. Vokan, Aru Islands.
Dr. Sclater (P. Z.
1875,
p.
87)
first
recognized the Aru Cassowary as a distinct
and gave a woodcut, showing a side view of the head and the wattles from
species,
iiont,
S.
accompanied by the iollowing diagnosis
" Casuarius similis C. australi ct crista iiari
in
:
modo
elevata
:
sed caruncula ceroids ana
media, ad apicem divisa.
"
Hab. Wokan,
"
Mus.
In
same volume our author
the
New
coast of
Aroensium [Beecuri).
inss.
Genoeusis."
civit.
Guinea, of which he believes that
that of the Aroo Islands,"
as
described as
also in the
Southern
mentions
also
C. sclateri,
namely
Cassowary from
it
However,
I
am
am
believe
species
it
has remained
form from
can be separated from that from the Aru Islands, where
it
Vokan group.
not able to give descriptions of well authenticated eggs of Beccari's Cassowary,
but there
named
same
of the
fully convinced that the
inhabits, so far as v^e know, only the northern island of the
I
is
south
the
This specimen was afterwards
beccarii.
but later on reunited with C. beccarii, and so
'Catalogue of Birds.'
New Guinea
C.
a
" probably
is
an enormous egg
C. bicaruncidatus,
is
in
the British
Museum
which, however, from
that of either C.
c.
beccarii or C.
c.
its
from the Aru Islands, which
large size
riolicoJUs.
It
I
am more
is
inclined to
measures 140
:
2'6
mm.
Its granulations are low.
Schalow
VOL. XV.
(/. c.)
— PART
V.
describes an egg brought from the
No.
2.
December, 1900.
Aru
Islands,
by C. Ribbe, measuring
t
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
nS
1'28
as
mm.
89
:
we know,
It
is
is,
however, not ascertained that
an egg of
it is
LSM. Casuarius
sp. S.
MiilL, Land- en Volkenkunde, p. 22 (Utanata, Princess Marianne Strait).
Casuarius heccarii (non
1875,
pi. Iviii. (S.
New
Mem.
ante:\,
Guinea)
1880. Casuarius beccarii (non
1881. Salvador!,
quod Aru!)
87,
p.
R. Ac. Sc. Torino,
1875,
(2)
sxxiv. p.
p. 484.
iii.
87
p.
1877. Casuarius australis (non Wall
x. p.
D'Albertis,
!)
19 (Fly River)
id., Ibis, p.
;
Casuarius sclateri Salvad., Ann.
Cornwallis Island, S.E.
1879. Sharpe, Ibis, p. 116 (type
New
now
;
'
This
as
occiput pale whitish blue.
i.
fig.
when
xii.
422
p.
id.,
;
Ann. Mus. Civ.
(coast opposite to
f.
Orn.
p.
Touan
or
300;
!).
laterally, very thin,
long and stout, black.
and
Wattles very
but in the old birds they get
and
t(n'ii
the thick scrub, so that often
This
of ragged ribbon-like appendages.
erect, as mucli as 6 feet
that of an
is
Count Salva-
Emu
[Droma'us).
high and the body being
Plumage
black,
Naked lower
head and
Hind-neck of
Fore-neck and nape brilliant indigo-blue.
a most intense bright scarlet.
sides of the neck, posterior
§ brilliant
from gape 130 to
^ bright blue. Total length about IGOO mm., bill
145
mm., tarsus 280 to 300 mm., inner claw SO to 95 mm.
140 and even
Juv. (two-thirds grown). Plumage dark brown, often nearly as black as adult,
scarlet, anterior
occasionally pale yellowish brown.
Fore-neck leaden blue
Hind-neck dull orange-yellow.
blue.
;
5
the Lirgest form of the C. casuarius group and also the
is
largest Cassowary, standing,
fully half as big again
pi.
!)
well illustrated in the series of iigurcs of D'Albertis's fine specimens in
monograph.
588
pp. 491,
Guinea only),
145;
p.
much compressed
Bill
number
they are very short or split into a
dori's
527,
p.
372.
either in fighting or wlien creeping through
damaged
New
A. B. Meyer, Journ.
Adult. Casque dull brown, very high and
large and long, sometimes over 7 inches,
:
Sydney Mail,'
preserved in Brit. Mus.
wjien fully adult curved over to one side.
Nuova Guinea,
D'Albertis,
:
Gen.
^NIus. Civ.
Guinea)
!)
197 (partim
New Guinea only
New Guinea only).
(partim
1895. Id., Cat. B. Brit. Mus. xxvii. p. 59G (partim:
Genova,
Proc. Zool. Soc. Lond.
Scl.,
;
Scl., P. Z. S.
1882. Id., Oni. Papuas. e Molucc.
1878.
evidently not the case.
is
D'Albertis's Cassowary.
Casuarilts casuarius sclateri Salvad.
•".
which,
beccarii,
c.
That the eggs of any
not the only Cassowary living in the Aru group.
Cassowary are more transparent than those of any other species
C.
Naked lower
sides
;
also
head and occiput pale
of neck, anterior ^ blue,
posterior f yellow.
Chick.
Head and neck dark
Longitudinal dorsal bands
with rufous, and
fiank extending
of dark
JIab.
down
Southern
variegated
with
black
bands
and
spots.
broad, about 1| inches wide, black, slightly variegated
A'ery
number
—
/.
e.,
a central one and two lateral pairs
;
one band on
the thigh on each side, and a second one indicated by a
brown irregular
iuid perliaps
I
5 in
rufous,
number
spots.
New Guinea
from Port Moresby
to
the Princess Marianne Strait,
even up to McCluer's Inlet.
have had several examples of
this
bird alive, though
none quite
adult.
They
—
irOX.
appear
be delicate and never to live long, and none ever seem
to
captivity
W. ROTHSCHILD OX THE GENUS CASUAEIUS.
\^
liich
species attains in a wild state.
tlie
are the largest Cassowaries
The
known.
110
to reach the size in
Adult wild specimens of
fine adult bird
this race
from which the descriptitm and
accompanying photograph were taken was shot by Mr. Emil Weiske on the Brown River
in S.E.
New
Guinea.
Fig. 1.
t'asiiariiii'
Sclater's
it
Cassowary was
cit.siiaritis
thought
at first
sclKfcri,
belong probably to Beccari's Cassowary;
to
was then separated, and afterwards united with the
original diagnosis of this form
" Medius, niger, casside
verticali,
brevissimo, crasso
interni breviuscula
"Long.
tot. 1'"
;
is
as follows
parum
loris et
;
latter
by Count Salvadori.
alta, crassa,
superne rotundata, latere posteriore lero
paleare unico, mediocri, ad apicem diviso
mento
'Ihe
:
violateis
;
;
ungue
digiti
capite et gula cseruleis, paleare rubro.
5U0, rostri hiat. 147 mm., tarsi 300, ungu. dig.
int.
80 mm."
T 2
HON. W. EOTHSCHILD ON THE GENUS CASUAEIUS.
120
A
most magnificent
and a number
of woodcuts of the heads and necks of
Genoa Museum and
the
was procured on the Fly River by D'Albertis,
series of this bird
them
are given in the
Annals of
Count Salvadori's splendid monograph of the genus
in
Casuarms, and a coloured figure, from a sketch of DWlbt-rtis, on plate
i.
(fig. 4)
of
the same work.
Some eggs from near Port Moresby, which
together with a chick, are
I received
covered with glossy glazed granulations, which are not at
more or
146
:
4.
1875
less
They are bright
separated from each other.
100, 146
:
and 146
!:»7,
Salvadori's Cassowary.
Casuakius casuarius salvadorii Oust.
?.
green, and measure 150
Casuarius tricarunculatus Beccari, Ann. Mns. Civ. Geneva,
1878. Salvad., Ann. Mns. Civ. Geneva,
1881. Salvad.,
Mem.
(2) xxxiv. p. 184
1882. Salvad., Ann. Mns. Civ. Genova,
xviii. p.
1882. Salvad., Orn. Pap. e Molucc.
p. 47.3
Salvad., Cat. B. Brit.
189(i.
Oustalet,
Mus.
Nouv. Arch. Mus.
iii.
Paris, (3)
717 (Warbusi)
vii. p.
413
;
;
;
591
xxvii. p.
XXIV.)
(Plate
;
263 (type in Mus.).
viii. p.
1878. (Feb. 23rd.) Casuarins salvadorii Oustalet, Bull. Assoc. Sc. de France, Bull. no. 539,
(" Warbu.si ")
Meyer, Jouru.
1881. Salvad.,
p.
Mem.
p.
350
;
1878. ScL, Proc. Zool. Son Loud. pp. 213, 214,
1879. Salvad., Ibis,
]03,
420;
pp. 419,
xii.
R. Ac. Sc. Torino,
18!)5.
:
mm.
95
:
connected, but stand
all
f.
Orn.
p.
202
;
Salvad.
fig.
213 (Wandarameu; head and ueck)
Ann. Mus.
Civ.
Genova,
xii. p.
;
A. B.
420;
105;
R. Ac. Sc. Torino, (2) xxxiv.
1882. Salvad., Orn. Papuas. e Molucc.
iii.
p.
488
p.
204,
pi.
(footuote)
i.
fig.
5
;
;
1895. Salvad., Cat. B. Brit. Mus. xxvii. p. 595.
1878. Casuarius altijugus Scl., Nature, xvii.
1879. Salvad., Ibis,
p.
A(hdt. Plumage black;
pointed.
scarlet.
;
casque high, brownish black;
and fore-neck deep blue.
inner half blue, rest dark
Chick.
375 (Wandammen)
long,
thick,
bill
and not
Head and upper part of hind-neck pale greenish blue, rest of hind-neck
Naked lower sides of neck blue anteriorly, purplish crimson pisteriorly.
Cliui, throat,
316 mm.,
p.
105.
bill
Wattles small and widely separated, base and
pink.
flesliy
Total length about
from gape 150 to 190 mm., inner claw about 80
Head and neck
rufous
;
rest of
H
m., tarsus
body o'eamy white or pale
longitudinal dorsal black bands variegated
with rufous.
800
to
mm.
The
buff,
with three
three lateral stripes
along the flanks and thighs are broken up iu'o a number of irregular brown spots
and patches.
Ilab. North-west
New
Guinea.
Casnarius casnarhis salvadorii was originally described by Oustalet and Sclater as
C.
salvadorii
Wandammen
and
C.
altijiir/iis
respectively,
from skins obtained by Laglaize near
on the south-west coast of Geelvink Bay,
in
North-west
New Guinea
;
;
121
HON. W. liOTHSCHILD ON THE GENUS CASUAEIUS.
and
Europe
but there are u certain number
Guinea, and which agree neither with Casunrius aisuariiis iVoni
of wild-shot Arfak birds
as
cominc from
New
Ceram, nor with
of birds sent alive to
;
from Aru, nor
C. c. beccarii
and these birds
in colour,
naked parts
nothinii; absolutely certain as to the colour of the
day we know
to this
I,
from Southern
C. c. sriateri
for the present, take
to
New Guinea
be Casmrius camarius salvadorii
as they agree in other respects.
The
bird from
which
I
made
the above given description lived two years at Tring
It is fully adult, and a male.
and one year in the Society's Gardens.
several younger ones, but none lived to maturity.
The
Genova,
Beccari in his ornithological letter (Ann. Mus.
shortly mentioned by
bird
as
vii.)
have had
I
agreeing in the form of the helmet with
C. casuarii(s,
but having
three wattles, namely, two lateral ones, and below these a smaller median third one,
was a young bird bought at Warbusi, a little to the south of Dorei. It was left at
Ternate to
eventuallv
grow up, where Laglaize saw
lost.
It
must, in
my
Oustalet says that the type of
Geelvink Bay.
it
still
alive
opinion, have been an
C. salvadorii
Warbusi, however,
is
came
in
July 1S77, but
aberration of C.
c.
it
was
srdvadoru.
also from Warbusi, at the bottom of
near Dorei, while
Wandammen
is
the place
southern end of the Bay whence the other specimens from Laglaize were said to
Therefore Salvadori (1881) thinks that Warbusi might be a mistake for
come.
A\andammen but I see no reason why it should not be correct, since C. tricaruaoaat the
;
latus
is
not a species according to
my judgment,
as
I
have a similar aberration of
Laglaize probably bought his specimens i'rom Bruijn or his hunters.
excited these birds extend the neck horizontally and emit five or six very
C. e. scIaUri.
When
several times,
loud grunting and grating sounds frequently, when very angry, they spit
of the hind-quarters, and peck and kick
ele\ate their wing-rudiments and the feathers
;
most
5.
viciously.
Casuarius CASUAEius INTENSUS Rothsch.
Blue-necked Cassowary.
K. O. Club,
1898. Casuarius casuarius intensus Rothscli., Bull.
viii.
(nos.
(Plate
Iviii., Ixiv.),
XXVII.)
j)1j.
.xxi, Iv.
dark brown ; bill pointed, short,
Adult. Casque very high and laterally compressed,
Plumage black. Size small. Occiput, nape, upper half of hind-neck, chin,
black.
brilliant .scarlet
and fore-neck deep brilliant blue. Lower half of liind-neck
Naked lower
black patch.
between the blue and red of the hind-neck an irregular
throat,
sides of
neck of an intense blue.
wattles on f .re-neck medium-sized, oval, close
Two
blue outer edge, and a few small spots pink.
together, but separated their entire length,
length of helmet 130 mm.
gape about 110 mm., tarsus 275 mm., greatest
Bin from
Young
(half-grown).
Plumage brown
;
head and neck dull blue
Lower naked sides of neck blue.
with
Young (three-fourths grown). Plumage br.,wn, mixed
;
lower hind-neck
dull orange-yellow.
Fore-neck
dark
blue;
hind-neck orange-red.
head
and
Lower naked
occiput
sides of
indigo-blue,
paler
neck bright blue.
a few black featliers.
round
ear.
Lower
HOX. W. EOTHSCHILD ON THE GENUS CASUAEIUS.
122
Young
(full-grown).
Plumage
deep blue, paler around the ear
black,
;
mixed with brown
feathers.
Head and neck
Naked lower
lower hind-neck scarlet.
neck
sides of
very bright blue.
Hah. unknown.
In addition to the type I liave had
five
others alive, and
they
all
agreed in having
the intense blue colour and the entirely blue naked lower sides of the neck
Cassowary remains
discovered.
" d ad.
more
erect
a very little-known form, and will do so
still
The original description is as follows
The casque differs from that of C. casuarius
till
;
but this
habitat
its
is
:
;
in
the wattles almost entirely blue instead of dark red
and neck uniform and very dark
;
much
being very high and
;
the orange of the hind-neck
the blue of the
much
lieat!
restricted
separated at the upper end from the blue by a black crescent-shaped patch
and
naked
;
sides of lower neck entirely uniform blue instead of red, bordered anteriorly with
blue."
Casuarius CASUAEIUS vioLicoLLis Rothsch. Violet-necked Cassowary. (Plate
6.
1899. Casuarius casuarius violicollis Rotliscli., Bull. B. O. C.
viii.
(uos.
lix., Ixiv.)
XXVI.)
pp. xxvii,
Iv.
Adult. Casque low, sloping back veiy mucli, laterally compressed, horny brown,
Bill very
green at base.
and pointed.
long, straight
Plumage
black.
Face bluish
Occiput and upper hind-neck pale greenish blue, throat and fore-neck brilliant
green.
Naked lower
Hind-neck orange-scarlet.
deep blue.
mauve, bordered anteriorly with a narrow
Two
of scarlet.
sides
of neck bright violaceous
line of blue, posteriorly with a
narrow
wattles on fore-neck large, round, and close together in female
;
line
small,
long, and wide apart in male, blue at base, rest fleshy pink.
Plumage brown;
Juv. (half-grown).
dull gamboge-yellow
Jm?).
;
neck and head dull leaden blue;
hind-neck
lower naked sides of neck dirty flesh-pink.
(three-fourths grown). Plunuige brown, with a few black feathers; liead
occiput pale dull blue
;
fore-neck dull indigo
;
hind-neck orange
;
and
lower naked sides
of neck dark pink streaked with purplish red.
Hab.
?
Trangan Island, Aru
The type
Islands.
of the species was imported along with two specimens of C. bicarunculatus
from Aru, and
is
a female, for she has laid an egg
but I have since received seven
;
others of this form, and they all agree in their characters.
As
the other islands of the
group are inhabited by different forms of Cassowaries, I
am
inclined to think that
Trangan Island must be the home of
I described this
1899
as follows
this interesting form.
form at the January meeting of the British Ornithologists' Club
in
:—
" This apparently undescribed form of Cassowary
is
most nearly
allied to Casuarius
casuarius salvadorii, but differs conspicuously in the colour of the naked parts and in
the very large
size,
which
fully equals that of C.
c.
anstralis.
;
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
much
" Bill
longer and straighter than in other Cassowaries.
123
Casque horny brown,
green at base.
Face and a broad band running down the side of the bill bluish green.
Base of lower mandible dark blue, with a yellow line running along one-third of the
length of the mandible on each side,
and
short, 3
X
2^ inches,
^^'attles at base of fore-neck very large, round,
blue at base, otherwise pink
])ale
whole length, but close together.
for their
Auricular
all over, entirely
orifice larger
Throat and fore-neck bright ultramarine-blue.
Cassowary.
neck pale greenish or eau-de-Nil blue.
Naked lower
Lower
separate
than in any other
Occiput and upper hind-
liind-neck brilliant orange-scarlet.
neck magenta-purple, bordered anteriorly with ultramarine-blue,
the magenta-purple space deeply carunculated and
sides of
posteriorly with orange-scarlet
;
sharply cut off from the red and blue borders, which are plain and smooth."
The " song " of this bird is an extraordinary performance, generally as follows
head and neck and remains in
It lowers its
out
stretched
straiglit
gradually inflating
head so that the
with great
its
bill
effort,
in
front
—
for about
and wattle clap together,
the skin of
its
shell
—with
head and neck
then,
;
the bill open and
bowing and jerking its
emits some barking grunts, apparently
neck being then so much inflated that the wrinkles
tlie
become quite obscure.
The type specimen laid an egg
but
a sound
it
position
—
seconds, \vith
fifteen
making
neck, without
this
:
last
which
year,
is,
needless to say, bright green
not properly developed, and a description of the surface and size
is
;
is
therefore better omitted.
7.
The Australian Cassowary.
Casuarius CASUAKius AUSTRALIS AYall.
(Plate
XXV.)
1854. Casuarius ausfraUs Wall, Ilhistr. Sydney Herald of June 3rd;
1857. Gould, Proc. Zool. Soc. Loud. pp. 2G9, 270
1858. G. R. Gray, Proc. Zool. Soc. Lend.
1860. Scl., Proc. Zool. Soc. Lond.
p.
1868.
Scl.,
p.
ii.
1866. Schleg., Zool. Gart. p. 180
Scl.,
iv,
;
196
310
Ibis, p.
;
360; Selileg. Jaarb.
p.
zool.
Genootsch. Nat. Art. ^lag.
200;
1865. Gould, Handb. B. Austr.
1867.
210;
p.
1862. Scl., Trans. Zool. Soc. Lond.
p.
;
Proc. Zool. Soc. Lond.
Lond.
p.
206
;
Scl. Proc. Zool. Soc.
Loud.
p.
557
212 (C jo/msom — aKsiral/x)
p.
;
;
Bennett, Proc. Zool. Soc.
473;
Proc. Zool. Soc. Lond.
p.
376
Ibis, p.
;
348;
1868. Ramsay, Proc. Zool. Soc. Lond. pp. 381, 388
1869. Gould, B. Austr., Sappl.
1870. Editors, Ibis, pp. 119, 120
1871. G. R. Gray,
(skeleton)
;
Ibis, p.
348;
;
Hand-list B.
;
71; Krefft,
pis. 70,
iii.
p.
2,
Scl., Proc. Zool. Soc.
1873. Sclileg., Mus. Pays-Bas, Strut/iiones,
Lond.
p.
85
;
9;
p.
1874. Ramsay, Proc. Zool. Soc. Lond. p. 325
1875. Scl., Proc. Zojl. Soc. Iioud.
no.
;
9851;
p.
547
Flower,
(in
Proc. Zool.
confinement);
Soc.
Lond.
p.
32
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
124
1876. Ramsay, Proc. Zool. Soc. Loncl. p. 119 (life-history)
i.
Rams., Proc. Linu. Soc. N. S. Wales
;
186;
p.
1877. Rams., Proc. Linn. Soc. N. S. Wales,
&
HartiiiEj, Ostr.
Mem.
1881. Salvad.,
Ostr. Farming,
R. Ac. Sc. Torino,
1882. Salvad., Oru. Pap. e AIolucc.
iii.
1888. Rams., Tabul. List Austr. B.
p.
p.
95;
473
&
Salv.
(2) xxxiv. p.
p.
376 (woodcuts of head)
pp. 196 (distribution),
ii.
327
Scl., Ibis, p.
pi.
192,
fig.
i.
;
;
3 (headj
;
;
19 (distribution)
Among Cannibals, pp. 98, 99;
Nests & Eggs Austr. B. p. 291, pL
;
1889. Lumholtz,
1890. North,
1891. Sharpe, Cat. OsteoL Spec.
1892. Schalow, Zeitschr.
1891. Schalow, Journ.
Coll. Surg.
xv. (egg)
iii.
439
p.
;
;
Ool. p. 11
f.
Orn.
f.
11.
p.
24;
1895. Salvad., Cat. B. Brit. Mus. xxvii.
594
p.
;
1896. Mcston, Proc. Roy. Soc. Queensland, x. pp. 59-6'J'.
1866.
Casuariiis jo/i/isonii F. Miiller, Australasian of Dec. 15th
pp. 241, 242)
1867. Carron
&
348 (maintains that Jofinsoni
Il)is, p.
1870. A. Newton, Ibis,
1873.
;
Krefft, Proc. Zool. Soc.
p.
is
;
correct name)
;
120 (contends with others that the name australis
Casuarius regulis (uomeu nudum) Rosenberg, Journ.
f.
Orn.
is
to be used).
390.
p.
Adult. Casque very high and massive, compressed hiterally, deep brown
and
stout, black.
;
throat,
Next
Plumage
casuarius.
blue
Lond.
;
1869. Krefft,
?
Proc. Zool. Soc. Lond. 1867,
;
Bennett, Proc. Zool. Soc. Loud. pp. 473, 474
482 Diggles, Orn. Austr. pi. & text
1868. A. Newton, Ibis, p. 348 (critical note on Diggles)
p.
{cf.
black.
lower hind-neck
bill
long
Sides of head, occiput, and upper hind-neck pale greeui-sli
scarlet,
running a
and fore-neck deep indigo-blue.
Two
blue, giving a purple hue.
;
casuarius sclateri, the largest of the forms of Casuarius
to C.
little into
the blue of upper portion.
Lower naked
sides of
wattles on fore-neck very
C'iiin,
neck mixed red and
large, deej)
pinkish red,
mottled with paler pink, blue at base.
Youug
Head and hind-neck
(quarter-grown).
fore-neck densely covered with short
downy
chestnut rufous.
Chin, throat, and
feathers of a brownish-buff colour.
Rest
of body clayey brownish yellow mixed with dark rufous.
Head and
Young (two-thirds grown). Plumage brown.
lower hind-neck orange
mixed with
Chick.
yellowish
;
fore-neck leaden blue
;
occiput dull pale blue
;
lower naked sides of neck blue,
livid purple.
Head and hind-neck pale rufous;
From base of neck to end
buff.
fore-neck yellowish
of
tail
longitudinal black bands, variegated with rufous,
an inch wide.
On
shoulder-girdle
down
the sides are three irregular
tlie
buff; rest of body
along the back run
three
broad
and each about seven-eighths of
wavy black bands extending from the
sides to the beginning of the metatarsus.
Hah. Northern Queensland.
I have kept the adtilt female, described above, since
eggs at Tring, and has
now
lived nearly
two years
1890
:
she laid some 40 or 50
in the Society's Gardens.
—
125
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
The first discovery of the fact that a Cassowary existed in Australia is due to
Mr. Thomas Wall, who, like so many other explorers, left his bones in the waterless
Wall procured a specimen, but this was lost, and
deserts of the Australian continent.
only a poor description, evidently made from memory, was published in the Illustrated
Sydney Herald' of June 3rd, 1854, by Thomas AVall's brother, William Sheridan Wall.
'
The account ran
as follows
:
was procured by Mr. Thomas Wall, naturalist to
the late expedition commanded by Mr. Kennedy. 'Iliis was shot near Cape York, in
one of tliose almost inaccessible gullies whicli abound in that part of the Australian
The head is
The Cassowary, when erect, stands about 5 feet high.
continent.
"The
first
specimen of
this bird
without feathers, but covered with a blue skin, and, like the Emu, is almost without
The body is thickly covered with dark brown wiry
wino-s, having mere rudiments.
On the head is a large protuberance or helmet of a bright red colour, and to
feathers.
blue and
the neck are attached, like bells, six or eight round fleshy balls of bright
The first, and indeed the
scarlet, which give the bird a very beautiful appearance.
only, specimen of the Australian Cassowary
was unfortunately
left at
Weymouth
Bay, and
Mr. Wall being most anxious for its preservation had secured
him to the spot Avhere, unfortunately for himself
it in a canvas bag and carried it with
well as in
and for science, it was lost. In the ravine where the bird was killed, as
in companies of
other deep valleys of that neighbourhood, they were seen running
has not been recovered.
On
seven or eight.
plentiful,
and
will
that part of the North-eastern coast, therefore, they are probably
be met with in
all
the deep gullies at the base of the high
hills.
The
was eaten, and was found to be delicious; a single leg aff"orded more
The
could dispose of at one meal.
substantial food than ten or twelve hungry men
in
the
strength
use of this
Cassowary possesses great strength in its legs, and makes
more strong and heavy than
same manner as the Emu. Their whole build is, however,
They are very wary, but tlieir presence may be easily detected
that of the latter bird.
which is taken up and echoed along the
by their utterance of a peculiarly loud note,
and it would be easy to kill them with a rifie."
aullies
doubt that it actually refers to
Most incorrect as this description is, there can be no
flesh of this bird
;
the Australian Cassowary, and
accept the
for
name
o£
austmUs
we can
iherefore, in accordance with all other authorities,
for this bird,
and need not substitute that oi jolumnii
it.
This form
The
is
most closely
allied to the
Ceram Cassowary and
to Sclater's Cassowary.
show the differences.
key, descriptitms, and plates, however, will
known only from Northern Queensland, from Cape York south to
acquainted with the lifeRockingham Bay. Of no other Cassowary are we so well
of the latter is that of Ramsay in the Proc.
history." The most complete description
This bird
is
Zool. Soc. Lond. 1870, pp. 119-123.
It runs as follows
:—
One of the chief objects of my visit to Rockingham Bay was
1900.
VOL. XV.— p.\rvT v. No. o.— December,
"
to
become acquainted
u
HON. W. EOTHSCHILD ON THE GENUS CASUARIUS.
126
In 1872
with the habits of this noble bird.
While
Brisbane on
in
specimen, the
bringing
first
alive to
it
arrived safely.
and, for the
I
first
my
had sent
collector,
Edward Spalding,
but with very poor results.
to this district for this purpose,
"
I
my way up
I
purchased by telegram a
tine
young
living
had been obtained and reared, and ultimately succeeded in
Sydney and shipping it to the Society, where I am glad to hear it
that
found also that several very young Cassowaries had been obtained,
This was great news
time, a nest and eggs liad been found.
;
and
I
I
was
most hospitably entertained and welcomed by Inspector Johnstone, who was the
first
need not relate how
to rediscover
I
made
haste to the Herbert River Police
all
Camp, where
and bring under the notice of others the existence of
remarkable
this
found Inspector Johnstone a true sportsman, as well as an ardent lover of
I am indebted to
nature, a zealous and energetic naturalist, and a careful observer.
this gentleman for much valuable information respecting the manners and customs
sj.ecies.
I
many
of the aborigines, and notes on the habits of
birds and animals
new
to
me, and
especially for information on the present species.
"The
Australian Cassowary
district of
is
a denizen of the dense dark scrubs scattered over the
Rockingham Bay, and extending
as far north as the
Endeavour River.
It
was tolerably plentiful only a few years ago even in the neighbourhood of Cardwell
but since the advent of sugar-planters, etc. on the Herbert River and adjacent creeks,
;
down and
these fine birds have been most ruthlessly shot
which
skins, several of
easily
enough procured
decreased, that
it is
can be obtained.
may be
I
saw used
I
ior hearth-rugs
destroyed for the sake of their
and door-mats.
Formerly they were
bnt latterly so wary have they become, and their numbers so
;
only with the greatest amount of patience that even a stray shot
know
of no bird so wary and timid
plentiful enough, and easily found in the soft
and although their fresh tracks
;
mud
on the sides of the creeks
or under their favourite feeding-trees, yet the birds themselves are seldom
now
seen.
most dense parts of the scrubs, wandering about
the sides of the watercourses and creeks, diving in through the bushes and vines at
Toward evening and early in the morning they usually visit
the slightest noise.
During the day they remain
their
favourite
in the
feeding-trees,
such
the native
as
figs,
Leichardt-trees,
and various
Acmena, Jamhosa, Lavidsonia, &c. they appear to be particularly fond of
the astringent fruit of the Leichardt-trees and of a species of Maranta, which produce
species oi
;
bunches of large seed-pods
filled
with juicy pulp, resembling in appearance the inside
of a ripe passion-fruit {Passijlora edulis).
sought after
;
the tame semiadult
biril
Fruits and berries of
which
I
all
kinds are eagerly
had the pleasure of forwarding
to the
Society (187-3) became so fond of the fruit of the Cape-Mulberry that he would allow
no one
to
come near the
tree
he had taken possession
devoured at a time as much as three quarts of loquats
'
and several
3 pounds).
I
found
tliat
in
This bird has frequently
(fruit of
Eriobotria japonica),
amount of bread per diem (about
the afternoons they frequently came out and
fair-sized oranges whole, besides its usual
In nature,
'
of.
127
HON. W. ROTHSCHILD ON THE GENUS CASUARIUS.
walked
alon^' the scrubs, or
along
tlie
quantities of pebbles and small rough-edged stones.
sweet potatoes
(in
and swallowed large
In confinement, plantains and
sides of tlie river or creeks,
while
large pieces, wliirh they swallow whole) are a favourite food,
cockroaches,
nothing seems to come amiss to them— grasslioppers, spiders, earthworms,
of their
flavour
tlie
They ascertain
caterpillars of all kinds, dough, and even raw meat.
and if not
by first taking it up in the tip of their bill and giving it a slight pinch
diet
;
throw it away.
invariably refused green loquats, but always picked them up in the
they
"I found
In confinement they become very tame, and may be allowed to
bill first to try them.
suitable, they
walk about the place without
restraint,
coming when
called, or
more often running
after
and following any one accustomed to feed them. If disappointed or teased, they not
or in
unfrequently show fight by bristling up their feathers, and kicking out sideways
witnessed on more
front with force sufficient to knock a strong man down— a feat I have
'
'
These birds are very powerful, and dangerous
than one occasion.
wounded,
to a tree
;
(^n
more than one occasion
a
wounded
the sharp nail of the inner toe
is
to
approach when
bird has caused a naturalist to take
a most dangerous weapon, quite equal to
as much execution.
the claw of a large Kangaroo, and capable of doing quite
" I found the Cassowaries to be excellent swimmers, and frequently tracked them
mile
On Hinchenbrook Island, situated about
across a good-sized creek or river.
and I have myself heard
the mainland, they have been frequently met with
U
from
them
;
calling at night
and early
in
of a least two miles from them.
the morning
as I passed
Mr. Johnstone informs
up the channel,
me he met
at a distance
with one swimming
his explorations while on the 'North-east
across a river of considerable width during
Their note, most usually emitted by the male, is a series
Coast Exploring Expedition.'
repeated, and continued for about three
of harsh guttural prolonged croakings quickly
the water at a distance of at
minutes f it is very loud, and may be detected across
resounding through the scrubs
three miles on a still night. I have listened to it
least
at a distance
it close and one of
of a mile and a half on land, and then thought
August and
noises I ever heard. They breed during the months of
the mo.t unearthly
September.
The
fir^t
black troopers, from
was found by some of Inspector Johnstone's
Mr. Miller, a settler on the Herbert River, purchased
nest procured
whom
One which he kindly
mentioned hereafter. The nest consists
some
of the eggs.
presented to
me
of a depression
is
of the light green variety
among
the fallen leaves and
scrubs is covered, with the addition of a few more
debris with which the ground in the
well concealed
The place selected is always in the most dense part, and
dry leaves.
the only two
eggs were five in number
by entangled masses of vegetation. The
from the
differed
and in both cases one of the eggs in each set
instances recorded
The others
colour, and having a much smoother shell.
others, bein- of a light green
patches of dark
covered rather sparingly with irregular raised
all have a rough shell,
smooth ground. In tlie pale (No. 1) variety
but bright green on a lighter green and
m
;
HON.
12S
AV.
EOTHSCHILD ON THE GENUS CASUARIUS.
in
both
thinly spread over the central portion than at the ends.
On
these raisings on the shell are closer together, and not so well developed
more
varieties they are
;
the whole they closely resemble the eggs of Casuarius bennettu, in which similar
variations are noticeable
in the middle.
am
I
;
but they are larger and of a greater diameter, being greatest
indebted to Inspector Robert Johnstone for the fine series of the
eggs of this species which at present grace
"The
species
my
collection.
following are measurements of some of the specimens of the eggs of both
:
" Casuarius austral is.
Width
Lengtli in inches.
"No.
1.
Liglit-green smooth
No.
2.
Dark-green rough
....
sliell
533
5-3
shell
in inches.
X
X
3v3
X
X
X
X
3"54
3-88
" Casuarius bennettii.
"No.
"
1.
....
Liglit-grcen smooth shell
5'65
No.
2.
Light-green rough shell
5-32
No.
3.
Light-green rough shell
5-34
No.
4.
Dark-green rough
5-2
shell
3-31
3-4
3'32
of Casuarius australis are of a dull rusty brown, the feathers having
The young
frequently a blackish shaft-stripe, giving to the back a streaked appearance.
first
year the plumage takes a deeper, lighter
brown hue, and black
appear mixed with brown, some being parti-coloured.
After the
feathers begin to
After the second season, at the
age of from 18 to 24 months, the black feathers predominate, and the helmet, which has
liitherto
been undeveloped, more like the shield of a Coot {Fulica), begins
keel or ridge in the centre, which rapidly increases in height.
on which
still
remain
a
The
to
show a
skin round the head,
few brownish hair-like feathers, begins to become wrinkled
and coloured, varying from bluish-greeu to orange on the lower part, and bright blue
on the sides of the neck, the wattles becoming carmine. The helmet still remains
comparatively small and undeveloped long after the wattles and naked parts of the
neck become coloured.
the fourth or
fiftli
I believe that
the helmet does not attain
is
carried
branches of trees striking the helmet slide over
greatly
impeded
jumping over
;
bordering the Herbert
but as
it is,
fallen trees
it
low
to the ground,
on to the back.
River and elsewhere, progress would
and louud
it
be
the Cassowaries traverse the scrubs with wonderful speed,
and logs wlien in the way.
A young
Society), while in the possession of Inspector Johnstone, during
its
and the vines and
Otherwise, in the
bird (the identical
specimen, I believe, forwarded by His Excellency the Marquis of
jumping out of
size until
year at least.
" In traversing these scrubs the head
dense vine-scrubs
its full
yard over a fence more than 6 feet in height.
6 feet 6 inches to the top rail,
on which
its
my
I
Normanby
visit
to the
succeeded in
measured the fence,
feet-marks were plainly visible
;
;
;;
;
;
;:
HON. W. EOTHSCHILD ON THE GENUS CASUAEIUS,
the length of the yard \\as only
moult
March
in
;
12x12
I
feet.
fouud
but the new feathers had uot
During these months specimens
tlie
made
all
times very spiteful, even attacking
At
share of their dislike.
to,
which
irritable
when
and frequently
unwell), and were at
very fond of bathing; the
tliey are
forwarded to the Society, was remarkable in
I
and might frequently be seen waiting at the
one came for water, when he would
May.
their appearance iu
but strangers chiefly come in for a
;
times I have noticed
all
semiadult bird before alluded
this respect,
keepers
tlieir
adult Cassowaries iu full
were remarkably
in confinement
sulky, even refusing their food (which they invariably do
129
pump
in the
yard until some
down
quietly under a copious shower, stretching
out his neck and ruffling his feathers up to allow the water to reach the skin. They
sit
do not like any exposure, and always endeavour
In the wild
to get out of the sun.
seldom leave the scrubs, and certainly never do so in the heat of the day
unless hard pressed but on the whole they are remarkably hardy, and bear confinement
state they
;
well.
In February last
purchased four
(187-")) I
which were obtained from some
young birds about G months
tine
the Herbert River district
settlers in
;
old,
these also
I
forwarded to England during the same month."
The eggs
of C.
c.
austraUs cannot be distinguished from
although most of them, but not
Eggs
are slightly larger.
all,
those
of C. casuarius,
laid
in
my
aviary
at Tring measure:— 140 93, 131:87, 136:88, 138:89, 130:93, 135:95, 132:95,
:
128
:
128 92, 129 92, 121
91,
:
:
:
91
mm.; two
in the British
Museum measure
130
:
84
and 143 91 mm.
:
8.
Casuakius bicarunculatus
The Two-wattled Cassowary.
Scl.
I860. Casuarius bicarunculatus Sclater, Proc. Zool. Soc. Loud. pp.
locality
186'2.
Id.,
uukiiowu)
(Plate
248, 249,
"211,
XXVllI.)
fig.
(youujr
;
Trans. Zool. Soc. Loml.
iv. p.
1862. Crisp, Proc. Zool. Soc. Loud.
p.
358,
fig. b, pi. Ixxiii.
137 (gall-bladder)
1862. Sclilegel, Jaarb. zool. Genootsoli. Nat. Art. Mag. p. 198;
1864. Id., Diereut., Vogels,
p.
239;
1866. Sol., Proc. Zool. Soc. Lond.
1866. Sclileg., Zool. Gart.
p.
p.
178;
168;
id.,
Nederl. Tijdschr. Dierk.
1867. Rosenberg, Reis naar Zuidoostereil.
p.
Scl.,
1872.
Scl., Proc. Zool.
Soc. Loud. pp. 150, 195,
pi.
xxvi.
1873. Garrod, Proc. Zool. Soc. Lond. pp. 470, 641 (anatomy)
1873. Schleg., Mus. Pays-Bas, Strutltiones,
1875. Rosenberg, Reist. Geelvinkb.
1878. Salvad.,
Ann. Mus.
1881. Gould, B.
fig. 1
Id.,
pp. 25U, .347;
Proc. Zool. Soc. Lond. p. 149 (young, alive);
1869.
1882.
lii.
52
New
p.
Civ. Genova,
Guinea,
v. pi.
p.
;
10;
117;
xii. p.
316
73; Salvad.,
(in
Bremen Mus.)
;
Rosenb. Malay. Arcli.
Mem.
R. Ac. Sc. Torino,
id.,
Orn. Pap. e Molucc.
(2)
(head)
Ann. Mus.
1894. Sclialow, Journ.
Civ.
f.
Genova,
Orn.
p.
xviii. p.
19 (error
:
414;
egg, Fly River
!
!)
p.
xxxiv. p. 185,
iii.
p.
475;
373
pi.
I.
HON.
180
EOTIISCHTLD ON THE GENUS CASUARIUS.
AV.
Mus.
1895. Salvad., Cat. B. Brit.
xxvii. p. 591.
1857. (?) Casuarius galeatus Wallace, Ann. Mag. Nat. Hist. xx.
1858. (?) C. emi'H (nou Latliam
18(51.
!)
(?) Casuarius kaiipi (non
p.
xxv. p. 252
lud.
134, quod unappendiculatus
p.
p.
(non
251) (Aru)
p.
iii.
1884. (?) Casuarius sp. A. B. Meyer, Zeitsehr. ges. (Jrn.
" Kudari,'" in the
Adult. Plumage black
deep, horn-brown.
Upper
On
p.
i.
the rest of
Young
(tliree-fourtlis
and head dull blue
;
p.
52.
Head and
is
occiput pale greenish blue.
scarlet
;
fore-neck dark blue.
a long pendent wattle, blue at base, deep pinkish red for
dull orange
;
fore-neck
wattles pale pink.
JTo^mr/ (Jialf-grown).
flesh-colour
Oru.
casque compressed at sides, not flattened at back, low and
Lower naked sides of neck deep blue.
grown)
Plumage brown. Hind-neck
suifece.
tlie
f.
296 (egg from Aru).
part of hind-neck deeper blue, rest of hind-neck
neck
Jouru.
Islands (fide Rosenberg)
Bill long, pointed, black.
eacli side of the
I
;
Aru
438;
p.
id.,
347.
p.
1867. Casvarins galeaius (non Vieiliot!), Rosenberg, Reis naar Zuidoostereil.
:
;
!).
1866. Casuarius aruensis Schleg., Nederl. Tijdsclir. Dierk.
Native name
!).
187 (sternum from Aiu).
Rosenberg!) G. R. Gray, Proc. Zool. Soc. Lond.
18G3. Rosenberg, N. Tijdsclir. Ned.
135 (non
(Aru Islands
p. -^77
G. R. Gray, Proc. Zool. Soc. Lond.
Hind-neck and wattles dull whitish
Plumage yellowish brown.
fore-neck and head dull greyish blue.
;
have received from Dr. Finsch, among a number of chicks of Cassowaries in the
Leyden Museum, one brought by von Rosenberg
group, which
difl'erent
is
clearly C. hicanmculatus.
from any other chicks
I
It
have seen
;
in
1S(")|J
from Kabroor Island, Aru
appears to be
rufous
it is
all
much
over,
faded, but
is
very
and has Ave dorsal
bands and two bands of irregular patches on the thighs of a darker, more brownish
rufcnis colour.
Wammer
Hah.
The two
and Kabroor, Aru Islands.
birds on
which the descriptions here given are based have been kept
by me, both at Tring and in
in
England
The
first
tlie
Society's Gardens,
and the older of the two has been
five years.
" Casuarius hicarunculatiis, a
recently obtained a
at
was the following note by
notice that appeared about this Cassowary
Dr. Sclater on a young individual without locality (P. Z.
Gardens
name
young example
Rotterdam.
It
is
S.
18G0, p. 211):
propose to apply to a Cassowary of which I have
I
for the Society in
easily distinguishable
exchange from the Zoological
by the throat-caruncles being
placed far apart on the sides of the throat, lighter colouring, &c.
shortly arrive in this country, I
new
species at the next
wattles were figured.
hope
to
the
Aiu
be able to give
Meeting of the Society."
Adult individuals were
Valentyn (Oud en nieuw Oost-Indien,
existed in
alive
Islands,
and that
iii.
it
p.
full particulars
This was done
first
(p. 249),
bird itself will
concerning
tliis
and the throat-
brought to London in 1872.
299)
differs
As the
first
from
mentioned that a Cassowary
C.
casuarius, but he gave no
HON. W. ROTHSCHILD ON THE GEKUS CASUAKIUS.
description of
is
not at
all
Wallace (1857)
it.
uncommon
in the
tells
Aru
us that a Cassowary (which
Islands, that
young
tlie
131
lie calls C.
galeatus)
numbers
are brought in
to
Dobbo, where they soon become tame, running about the streets, and picking up all
It is probable that the birds he saw were mostly C. hicarunsorts of refuse food.
A
culatus.
sternum only was procured by Wallace.
Our first knowledge of the
who (' Reis naar Zuidoostereil.')
actual
home
of this bird
is
to
von Rosenberg,
Aru
Islands on April
due
actually shot a Cassowary on the
He
15th, 1SG5, which he called C. yaleatus, but which pro\ed to be C. biranmcnlatus.
the
(from native sources) that the pairing-season is in June and July, and that
tells
us
sexes live separate exce])t at that time of the year
rt)ugh nest and sits on the eggs for about 28 days
:
;
that the female
makes a
sort of
but both these latter reports are
proved that
doubtless erroneous, as the observations in the Zooloj;ical Gardens have
longer.
much
the male alone which sits, and that the time of incubation is
it is
Rosenberg
also tells us that not
more than
five
some more eggs are placed outside the nest
This ridiculous story— which is also
hatched
!
to
serve as food
for the
young when
Ostrich— is, it is needless to
of his want of knowledge of the
told of the
In spite
former assertions.
say, just as reliable as the
eggs are found in the nest, and that
discovering the
habits of the birds, however, Rosenberg deserves full credit for
home
Islands, Kabroor and
of C. bicanin culatus, which he obtained in the northern Aru
"
Kudari." Specimens have from
native name of this bird in Aru is
Wammer.
The
time to time arrived alive in Euroi-e,
aiul
have now two
1
alive,
both received when
almost chicks.
Eggs from the Aru Islands have been described
as those of C. bicarunculutus, but as
jjriori any
two or three Cassowaries are found in that group of islands, there is not a
not seem to differ very
absolute certainty about any of these eggs, which, however, do
hicaruaremarkably from those of V. casuarius and its aUies. The egg identified as €.
culatus in Nehrkoru's collection
from
the Fly
River, but in
is
said by
Nehrkoru's
'
Schalow (Journ.
f.
Orn. 1894,
Katalog der Eiersammlung
'
p. lU) to
be
said
to
it
is
Nehrkorn says
It is iherefore not an egg to depend on.
be from the Aru [slands
eggs of Dronueus, and that it
that it is so dark blue-green that it resembles the
According to Schalow its glazed granulations are less
measures 133 :9U ram.
but this may be an individual
continuous and less numerous than th.jse of C. casuarms,
" durchSchalow also distinguishes between Cassowary eggs that aie
character.
"
the light; but this is by no
scheinend" and " undurchscheinend when held against
are emptied,
difference, being mostly due to the way in which the eggs
!
means a
!
specific
and the thickness of the egg-shell varies also considerably.
Rosenberg, have more
Four eggs in the Leyden Museum from Kabroor, collected by
The corrugations
agree in other respects.
or less lost" their original green colour, but
The measurements are :—
elevated.
are more or less interrupted and more or less
129:87, 132:87, 121:82, 129:80 mm.
HON. W. EOTHSCniLD ON THE GENUS CASUAEIUS.
132
Casuaeius unappendiculatus Blyth.i
9.
One-wattled Cassowary.
(Plates
XXIX. &
XXX.)
1860. Casuarius
sp., Blytli,
Hist. ser. 3, vol.
112
vol. xxix. p.
Ibis,
145)
vi. p.
(juv.,
193, aud Sclater, P. Z. S. p. 210 (rcpriut Aim.
p.
no
307
p.
;
unappendiadatus Blyth, Journ. As. Spc. Beng.
Sclileg., Jaarb.
;
cum
p. 239,
1866.
Scl.,
34
iii.
vol. xxs. p. 103.
pis. 74,
75
iv. p.
p.
coast)
533 (exact
p.
locality,
1878. Salvad., Ann. Mus. Civ. Genova,
= uniappend.,
woodcut)
R. Ac. Sc. Tor.
1882. Salvad., Ann. Mus. Civ. Gen.
53
Gray, Hand-list,
;
Threshold Bay)
346
xii. p.
597
p.
(3) viii. p.
Tijdsclir.
iii.
9852
p. 2, no.
;
218
417, note;
Scl., Ibis, p.
;
p.
117
;
;
;
Rosenb., Mai. Arch.
p.
396
ii.
id.,
fig.
6;
Orn. Pap. e Mol.
iii.
p.
490;
264.
Mag.
1863. Rosenb., Natuurk. Tijdschr. Ned. Ind. xxv.
1866.
Ned.
Itid.
xxiii.
43,
p.
pi.; id.,
Journ.
t.
Orn.
i.
1862. Schleg., Jaarb. zool. Genootsch. Nat. Art.
1864. Id., Journ.
id.,
(egg)
ii.
Rosenb., Natuurk. Tijdschr. Nedeid.
p. 44, pi.
;
;
3, pi.
kaiipi
p.
(chick, Sorong)
414, no. 5;
1895. Salvad., Cat. B. Brit. Mus. xxvii.
1896. Oust., Nouv. Arch. Mus. Paris,
179
Rosenberg, Reist. Geelviukb.
;
xxxiv. p. 205, pi.
(2)
xviii. p.
1893. A. B. Meyer, Abh. Mus. Dresd. no.
C
Diereutuin,
;
1886. Beddard, P. Z. S. p. 145 (anatomy)
1861.
mala,
(juv., fig.
id.,
10;
1877. Forbes, P. Z. S. pp. 307, 313, 314, 316 (bursa Fabr.)
Mem.
74
;
1874. A. B. Meyer, Sitzb. der k. Ak. Wiss. Wieu, vol. Ixix.
1881. Salvad.,
pi.
198, pi.;
;
1873. Sclileg., IMus. Bays-Bas, Struthiones,
{kaupi
359,
p.
Zool. Gart. p.
Sclileg.,
;
250 (Salwatti and opp.
p.
1871. Schleg., Nederl. Tijdschr. Dierk. vol.
P. Z. S.
iv. p.
Genootsch. Nat. Art. Mag.
zool.
descr. of egg)
(first
1869. Gould, Suppl. B. Austr.
Scl.,
113
fig.
P. Z. S, p.
Dierk. vol.
1875.
ser. 3, vol. vii. p.
C. uniappendiculatus, Bennett, Ibis, p. 403,
1862. C. vntappendiculatus Sclater, Trans. Zool. Soc. Lond. vol.
typus)
Nat.
(head and neck, from live bird in Amsterdam).
pi. xiv.
C
Ann. & Mag. Nat. Hist.
locality), reprint
C. unoappendiculatus, Blyth, Ibis,
1861.
& Mag.
Casuarius unappendiculatus, Blytli, Journ. As. Soc. Beng.
;
f.
Orn.
Scl., P. Z. S. p.
p.
168
{
p.
p.
251
199;
;
136
= umuppendiculatus)
1867. Rosenb., Reis naar Zuidoostereil.
p.
;
Schleg., Zool. Gart. p. 180;
52
1873. Schleg., Mus. Pays-Bas, Struthiones, p. 12
;
Rosenb., Journ.
f.
Orn.
p.
390
(definite assertion
of identity with C. uniappendiculatus)
1894.
C.
unoappendiculatus Schalow, Journ.
Native names:
"Mambeba"
Adult. Casque low, very
Head and upper
'
By an
f.
Orn.
at Sorong, " Kallo
much
p.
23 (egg).
" in Salwatti {Rosenberg).
flattened behind, black; bill short, thick, and black.
part of hind-neck pale blue, with a greenish tinge.
Cheeks dark
error, on the four Plates of the various forms of C. unappendiculatus, as well as on the distribution
map, the name
is
spelt uniappendiculatus.
The
original spelling, however,
is
without the
i.
— W.
11.
—
•
;
HOX. W. ROTHSCHILD ON THE OEXUS CASUARIUS.
133
blue;
face-wattles louii', broail and thin, very bright dark blue.
Chin dark blue;
throat and a small part of hind-neck dark dull olive-yellow; fore-neck and rest of
A
hind-neck dark golden-yellow.
golden-yellow transverse patch on occiput.
lower sides of neck dark orange-yellow.
Height from casque
very large.
when
8 inches
ground about 5
to
;
Naked
wattle dirty mauve.
Size
when walking,
6 feet
feet 6 inches
erect.
Total length about 1G55 mm.,
(After Salvadori,
Young
Plumage black
I.
from gape 133, tarsus 280, claw of inner toe 84.
bill
c.)
Casque brown
(half-grown).
plumage yellowish brown neck and naked
Head, chin, and occiput dirty bluish
;
;
lower sides of neck pale yellowish flesh-colour.
green
legs yellowish.
;
Head and neck uniform rufous; throat buff; body creamy buff, with five
brown longitudinal bands on the body and a line of broken brown patches along
Chick.
distinct
the thighs.
N.W. New Guinea
Hah. Salwatty and opposite coast of
Ram, Sorong).
The first communication
editorial note in 'The Ibis'
C'alcutta, Jan. 8th
of the
Common
its
much
name
the
:
'
and a long
(first)
plumage,
size
of the legs,
Journal of the
it
it
naked yellow skin down each
of a
is
size,
much
easy to perceive that
is
in the
side of the
lighter colour than the
young
two of which are kept along with
Society
Asiatic
of Bengal,'
nobis,
n.
s.,
from
pendulous caruncle in front of the neck.
much paler
common C.
same age of the
the followiuf
'
when full-grown
Blyth
first
—
half-grown, and
it
is
in his last letters (dated
it
it
is
a
"
" Casuurius unappcndicidatus,
single
strip of
Cassowary of the same
smaller species.'
In
— " Mr. Blyth
:
present
and from the
for 1860, p. 193:
speaks of an apparently new species of Cassowary (Casuanus)
Babu Rajendra Mullick of Calcutta
It has a yellow throat, a
single yellow throat-wattle,
In
referring to the One-wattled Cassowary
& 21st)
in the aviary of the
neck.
(Threshold Bay, Tangion-
same paddock.
in
tlie
colouring of
its
its
peculiarity of
published
the
having but a
Specimen apparently more than
plumage than specimens of the
galeatus, two fine examples of which are associated with
In lieu of
tlie
two bright red caruncles of the
latter, the
new
species has but a single small oblong or elongate oval yellow caruncle, and the bright
colours of the naked portion of the neck are differently disposed.
throat are smalt-blue, below which
is
The cheeks and
a large wrinkled yellow space iu front of the
neck, terminating in front in the oval button-like caruncle, and
its
lower portion being
continued round behind, while on the sides of the neck the yellow naked portion
continued
dowu
to its base, the bordering feathers
this lateral stripe of
skin
is
unfeathered skin
;
— PAET
V.
No.
4.
or less covering and concealing
on the hind part of the neck the bare yellow
not tumous and corrugated as in the
VOL. XV.
mure
is
December, 1900.
Common
Cassowary, where also this part
x
.
HON. W. EOTHSCHILD ON THE GENUS CASUARIUS.
134
is
The casque is about equally developed
the new species are smaller, from which
The
legs of
the two species.
at this age in
bright red.
I
doubt
if it
attains to quite so
large a size as the other."
The next we hear concerning this species
this grew
Zoological Garden of Amsterdam
;
home,
for
the arrival of an immature bird in the
up, and thus was afterwards the
Also in this case Rosenberg was
described and figured by Schlegel.
discover the actual
is
he obtained
fii'st
the
adult
first
to
on the island of Salwatti, and afterwards
it
specimens became known from several places on the opposite coast.
The
Amsterdam
bird in
laid a pale
dark green, and measuring 136
my
have an egg in
I
who had
1871,
it
in
89
:
green egg, "thickly covered with raised spots of
mm."
collection (from the
Buckley
exchange from ''Mr. Franks":
from the Amsterdam dealer Frank, who had
on
when
"
label
its
Maart 1865
The
it
this
means probably that
from the Zoological Gardens,
(or 6, tlie 5 being obliterated),
This egg
bought from T. Cooke,
for
it
was
has
being about the time
tliis
from other Cassowary-eggs
differs
it
in
having
round knobs, like pin-heads, few of them being connected
single glazed
with each other.
{I.
"
the bird laid eggs there).
numerons
collection,
descriptions of the two eggs of C. unappendiculatus by Schalow
c), both from different sources, agree with mine
— the peculiar
isolated piu-head-like
elevations being well described and discussed as distinguishing the unajipendiculatus-
egg from
there
is
One
all others.
no doubt, as
it is
of those in the British
Museum, about
the origin of which
one of the eggs laid in Amsterdam, agrees also with these in
having the separate knobs, while two others, from Frank, and one said to have come
from Salwatti, are more like those of Casuarius casuarius.
knobs measures 137
:
88 mm.,
while Sclater gives only 136
151
:
81, Schalow's 146
said
like
measure 145
all
It
:
90,
145
in
96, 148-5
:
49'5 mm.,
On
Sclater also speaks of raised "spots" only.
89.
the
:
the
91,
145
:
90,
143
:
91,
147
:
92, 142
must therefore be
left to
95,
145
:
96
mm.
They
future research whether the peculiarly isolated knobs on
C.
unappendiculatus are a specific character
present this view cannot be taken, unless the identity of
Museum and
As
:
very large.
some well-authenticated eggs oi
usual,
Kosenberg
of some of those in the British
Museum
we know nothing definite respecting
tells
got angry
it
is
all
;
but at
the eggs in the Leyden
doubted.
the biology of this bird in a wild state.
us that an immature bird he got in Ternate was running about freely
and was much attached
it
:
the isolated
Leydeu Museum, sent by Bernstein and Rosenberg, and
to be from Sorong and Salwatti, do not show these characteristic isolated pin-headThey
elevations, but rather a more connected network of glazed granulations.
other hand, seven eggs
are
:
my own
The one with
put up
to people,
its
while
it
was a
fierce
enemy
to cats
and dogs.
When
feathers and emitted a peculiar blowing cry, followed by a
grunting like that of a young pig.
HON. W. ROTHSCHILD OX THE GENUS CASFARIUS.
10. Casuarius
wary.
1875.
J obi Island
Cdsiiarhts pajjuanus Rosenberg, Rcist. Occlvinkb. p. IIT (.lobi)
Ann. Mus.
Civ.
Gen.
vii. p.
1875. Casuarius occipitalis Salvad.,
1876. Scl., Ibis,
p.
245 (note)
300
pp. 203,
Mem.
(footnote)
;
;
xii.
p. 4.23;
Scl., Ibis, p.
1882. Salvad., Ann. Mns. Civ. Gen.
1893. Meyer, Abh. .Mus. Dresd. no.
f.
Orn.
(2) xxxiv. p.
xviii. p.
22 (egg)
p.
ii.
fig.
pi. ii. tig.
id.,
;
7 (head)
598
p.
494
;
;
viii. p.
" Orawei," Jobi {Rosenher[i)
deep broun.
iii.
;
204,
pi. xiv.
;
f.
Ool. p. 19.
" Orawai " at Ansus on Jobi Island [A. B. Mei/er).
Adult. Casque very high, coniral, flattened posteriorly.
Iris
;
Orn. Pap. e ^h,].
4 (egg)
1892. Casuarius westermatmi lley (nee Sclat.), Zeitselir.
:
Orn.
1'.
;
p.
1896. Oust, Nouv. Arch. Mus. Paris, (3)
209,
414, uo. G
3, p. 29, pi.
1895. Salvad., Cat. B. Brit. Mus. xxvii.
Native names
482; A. B. Meyer, Journ.
;
R. Ac. So. Tor.
1894. SchalovT, Journ.
J obi Beccaii,
Cusoitr di
;
718.
il)id.
1878. Salvad., Ann. Mus. Civ. Gen.
1881. Salvad.,
Onc-wattled Casso-
XXXI.)
(Plate
(?)
unappendiculatus occipitalis Salvad.
1 P.o
Beak long and
very stout.
Face, head, chin, throat, and upper half of neck deep blue, rather
Face-wattles long, thin, pendent, and much swollen at
Lower half of neck deep yellow or pale orange.
Pear-shaped wattle
yellow, more or less suff'used with blue.
Naked sides of lower neck deep crimson.
paler on hinder part of neck.
their base.
On
the occiput
is
a transverse patch of dull dirty orange.
Plumage black and coarse
;
legs brownish grey
;
bill
and casque black
in
most
fully-
adult specimens, but some occasionally retain the greenish-horn coloured or pale olive
helmet of the immature bird.
The measurements
about If m.,
bill
Legs dark
olive.
of the type {cf. Salvadori, I.e.) are as follows:
from gape 137 mm., tarsus 2S0 mm.,
YotiiKj (full-grown).
Plumage brown
green where deep blue in adult
;
face-wattles absent
;
—Total
nail of inner toe
;
70
length
mm.
miked parts dull bluish
dull yellow instead of pale orange on
lower half of
Casque dull green.
neck, and dirty red mixed with yellow on lower sides of neck.
Yomifi (half-grown) similar to that of C. unajjpendiculatus, but casque bright green.
Chick.
Head and hind-neck
pale rufous; body pale buff; back with five black bands
variegated with rufous; flank and thigh
This form
its
differs
from
C.
stri])es
broken up into irregular
unappendiculatus principally in
its
spots.
more slender build and
greater height, the greater extension of blue on the neck, the crimson lower sides
of the neck, and the very high casque.
C.
It inhabits the island of Jobi
and perhaps also the opposite coast of Geelvink Bay.
There occasionally occurs
this form,
in
and probably also
in
all
the other forms of
unappendiculatus, a curious melanistic variety, where nearly the whole neck
and only a small spot of yellow remains
at
the base of the hind-neck.
x2
is
blue
This was
HON. W. EOTHSCHILD ON THE GENUS CASUARIUS.
136
name
described by Oustalet as a distinct species under the
may stand
of Cnsuarius laglaizei
;
it
as
Casuarius dnappendiculatus occipitalis aberr. laglaizei.
[1893. Casuarius laylaizei Oust., Bull. Soc. Philom.
Oust. Nouv. Arch. Mus. Paris,
purchased by Laglaize
type
;
and measurements and plate
do not believe
It
is
to
p.
Museum.
Paris
show the supposed
(Jobi Islaud)
;
1890.
(Specimen killed on Jobi and
pi. xv.
iu
C. laglaizei,
no. 9, pp. 1-3
v.
(8)
265,
Careful description
specific value, in which,
however, I
at all.)]
most probable that
Ak. Wiss.
(Sitzber.
(3)
of
viii.
Ixix.
was
it
p.
C.
imaj^pendiculatus occipitalis of which A. B.
217) says that
occurs on Jobi, where
it
Meyer
called
is
it
"Orawai," and of which Rosenberg had received some reports from the natives when
stating
('
Reistochten,' p. 117,
occurred also on Jappen
— the
and Malay. Arch.
usual Dutch
p.
name
563) that Casuarius ]}apuanus
for Jobi
Island in Geelvink Bay.
Beccari brought a grand adult male from Ansus, on Jobi Island, to Italy, and this was
at once recognized
by Count Salvadori as an undescribed form.
to Beccari's interesting ornithological letter in the
our learned friend diagnosed this form as follows
:
In a footnote appended
Ann. Mus. Civ. Geneva,
vii.
p. 718,
—
" C.\suABius occipitalis mihi.
" Casside
compressa, postice
depressa
appendice
;
capite, gula et parte superiore cervicis caeruleis
inferiore flavis; area
more
nuda
'
macula
laterali colli imi carnea."
detailed Italian description.
close relationship this bird
;
had
Catalogue of Birds,' xxvii.
ut videtur, nulla
occipitali et colli parte
This diagnosis
is
nuda
followed by a
Needless to say, the Count always recognized the
to Casuarius
p. 599,
colli antici,
unappeiidiculatus from Salwatti.
he says that
it is
In the
" very similar to C. uniappendi-
culatus, but differs in having a triangular yellow patch
on the occiput, the posterior
surface of the casque narrower, the single caruncle smaller."
:
1
unless
more than one occur on that
thus closely resembling those
It
—
The measurements of the type are, according to Salvadori
Total length about
m. 670 mm., bill from gape 137 mm., tarsus 280 mm., claw of inner toe 70 mm.
One egg obtained by Doherty on Jobi Islaud, and doubtless belonging to this form,
measures 144
1 1
.
wary,
:
95
island,
shows a gcjod many rather isolated knobs,
of C. uiiappendiculatus,
which show
Casuarius unappe>.dicul.\tus aurantiacus Rothsch.
(Plate
this
character.
mm.
Eastern One-wattled Casso-
XXXII.)
1899. Casuarius uniappendiculatus aurantiacus Rothsch., Bull. B. O. C. vol.
p. Ivi (21st
June)
Adult. Face, head,
;
id.,
cliin,
P. Z. S. 1899, p.
774 (June 20th
;
viii.
published Oct.
p.
1
(17th May),
1st).
and nape of neck bright turquoise-blue, darker round eyes
—
HON. W. EOTHSCHILD ON THE GENUS CASUAEIUS.
and on
Rest of fore- and hind-neck and naked lower sides of neck deep
face-wattles.
bright orange, without any red.
On
Throat- wattle blue.
large transverse patch of bright orange.
the occiput a very wide and
Face-wattles large and very thick, balloou-
shaped when inflated by the bird during anger.
compressed
137
and thick.
Bill short
and not flattened posteriorly or hardly
laterally,
coarse; legs brownish grey;
Build slender and
pale brown.
iris
Casque low,
Pkimage black and
so.
tall,
but not so
tall as
either C. unappendiculatus or C. u. occipitalis.
Young and
chick unknown.
This form
differs
both from
C.
unappendiculatus and
C. u. occipitalis in its
slender
build and paler blue colouring, but principally in the uniform orange of the neck, the
compressed and posteriorly not flattened casque, and in the thick swollen
face-wattles, which are very similar to those of the following species, Casuarius philipi.
laterally
Hub. German
This form
is
New
Guinea.
known from one individual living at present in the Zoological Garden
Keulemans made the accompanying picture. It is most interesting
of Berlin, where Mr,
to find the one-wattled
group of Cassowaries extending
have no doubt whatever that
My
original description
my
aurccntiacuH
{I. c.) is
as follows
" Face, cheeks, and occiput pale sky-blue
to
Kaiser-Wilhelmsland, and
a very good subspecies.
is
:
;
throat dark blue.
Occipital patcii, fore-
neck, hind-neck, and lower sides of the neck deep reddish orange.
green, and
much more compressed
laterally
I
than in
C.
Casque horny
unappendiculatxis.
Long cheek-
wattles absent, but the sides of the face distended as in C. philipi."
12. Casuarius unappendiculatus rufotlnctus, subsp. nov.
Red-tinted One-wattled
Cassowary.
Casque greenish horn-colour, compressed
plumage black, mixed with brown,
otherwise in full colour.
on the occiput
is
Head,
as the bird
face, occiput,
laterally
is
scarlet,
changing
sides of
it is
to
about 1 inch wide.
;
Chin and throat deep indigo-blue,
which colour runs down the centre of the fore-neck
the wattle,
;
not quite in full adult plumage, though
and upper hind-neck bright pale blue
a large patch of orange-brown.
where
and not depressed posteriorly
in a triangular banil to the base of
Wattle blue.
Sides of fore-neck orange-
crimson when excited and the throat distended.
Lower naked
neck deep crimson, with a narrow anterior yellow border.
Hub. unknown.
I
had
this bird as a
but since
is
it
chick in striped plumage, and
it
was reared
has been in this Society's Gardens (15 months)
very different from any other form of C. unapjpendiculat us.
it
as a pet at Tring,
has come into colour and
It
is
now
3 years old.
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
138
13.
Sclater's Cassowary.
Casuarius philipi Rothsch.
1898. Casuarius
Almost
Rothsch., Nov. Zool.
jtliUipi
brown behind, darker
Plumasc black
;
front
in
much
feathers long and
same time depressed
and apparently rather
short.
webs being
curled on the body, the
close
other Cassowaries, more like the tip of a freshly-moulted
and not disintegrated as
in
feather of Dromceus.
Feathers of the
of the tail-feathers
at the
the colour of the casque pale
;
thick
bill
;
and
laterally
thus combining both forms of casque
posteriorly,
whitish
418.
v. p.
Casque very high, compressed
rtfJiilf.
XXXIIl.)
(Plate
rump and
when fresh-moulted
enormously lengthened, some
tail
Head, occiput,
trailing on the ground.
and upper part of hind-neck pale greenish Nile-blue
face,
fore-neck and uppersides of
;
lower hind-neck lemon-yellow, flecked
neck and sides of nape deep indigo-blue
Lower naked sides of neck bright cherrywith a few reddish spots on the edges.
;
when extended and
Cheek-pouches,
crimson.
lavender-blue
very thick
;
The
colour.
feathers
of
on the lower part of the fore-neck
two-thirds cherry-red, lower third blue.
clay-brown.
Cry unlike
all
On
a single round
is
of
dark
a
come very high up and
neck
the
during anger,
inflated
the occiput
flat
are
wattle, upper
a broad patch of dull
is
the other Cassowaries, being very loud and resembling
a deep roar.
the
quite
This,
colour of
its
naked
ordinary shape.
It
most remarkable Cassowary,
parts,
is
its
curious casque
New
only
it
Zealand Dinornis
to the
giant
a
is
by
distinguished
and feathers, but
every other respect
be compared with the extinct
not
short from the ground
quite as
Casuarius benveUi, but in
is
;
also
by
fact,
{Pacliyoriiis)
it
extra-
its
back as
level of its
in
tlie
can only
elephantojyus as
regards proportions.
Hah. unfortunately unknown.
It
was shipped
to
London irom
Calcutta, and
when
it
first
came
to
the Society's
Gardens was in brown plumage, and the colours of the head and neck were not
It was thought to be a young C. unappendimdatus, but as soon as it began
developed.
to
all
come
into colour, I
saw
it
was something quite new,
in fact
it is
the most distinct of
the species of Cassowary.
1 originally described
follows
this bird
when
it
was much younger.
My
description was as
:
" This
new
species is founded
London, which, though
far
on a bird now living in the Zoological Gardens,
from adult, being almost brown in plumage, appears
full-grown, and the naked parts are fully coloured.
appewliculatiis, but differs
much
to
be
C. uni-
in colour.
" Plumage when adult evidently black.
horn-colour.
It is closely allied
to
Casque
as yet undeveloped, pale yellowish
Throat and fore-neck deep purplish blue.
A
single
small wattle on
;
HON.
fore-neck, round and
;
EOTHSCHILD ON THE GENUS OASUARIUS.
\V.
not pear-shaped as in C. imiajypendiculatus
flat,
wattle purplish red, rest dark blue.
high up the neck.
upper third of
;
Hairy feathers of neck very thick and
reacliin-;
Head, occiput, and upper half of hind-neck very pale greenish blue
lower half of hind-neck pale orange-yellow.
Naked
crim?on, fading into cherry-red on the edges.
very low on
139
the
;
skin on lower sides of neck deej)
Legs very stout and short
body
;
set
and very bulky, giving the bird the exact shape Dhwrnis
legs
elephantopus must have had.
" Ilah. Probably Eastern German
"
Named
in
New Guinea ^
honour of Dr. Philip Lutley
PAPUANUS Schleg.
14. CASUArxius
Sclater."
Westerinann's Cassowary.
(Plate
1871. Casnarius heiinetti (uou Gould, 1857) Sclileg., Nederl. Tijdsclir. Dierk.
1871. Casuarius pcipuunus Sciileg. (es Roseubei'g MS.),
1873. Schleg., Mus. Pays-Bas, Struthiones,
p. 11
(Andai)
1875. Roseub., Reist. Geelvinkb. pp. 81, 117,111,
(Andai)
pi.
17
;
51
c. p.
;
Roseul)., .Jourii.
;
53.
iv. p.
Salvad. Aim.
f.
Oni.
p. ."390;
Gen.
^Iiis. Civ.
vii. p.
717
;
1878. Rosenberg, Malay. Arcliipel,
1879. Salvad.,
t.
XXXIV.)
Mem.
563; A. B. Meyer, Jouru. L Orn.
p.
R. Ac. Sc. Tor.
1882. Salvad., Ann. Mus. Civ. Gen.
1894. Schalow, Jonru.
f.
Orn.
p.
xxxiv. p. 210,
(2)
4li, no. 7
xviii. p.
20
id.,
;
Oni.
pi. ix.
1895. Salvad., Cat. B. Brit. Mus. xxvii.
1896. Oust., Nouv. Arch. Mus.
J*ap. e
203;
;
Mol.
iii.
p.
I'J.J
:
(egg).
1871. Casuarius kaupi Sclat. (nou Rosenberg, 1861), P. Z. S.
1873. Sclat., P. Z. S. pp. 147-150,
pp. 200,
8 (head)
pi. ii. fig.
(Mansiuam)
627
p.
(.Maiisliiara)
;
;
599;
p.
265.
(3) viii. p.
1872. Casuarius papuensis Sclater, P. Z. S. pp. 119, 150.
1874. Casuarius westennanni
1875. Scl., P. Z. S. pp. 85, 87, 380,
1877. Gould, B.
New
Guinea,
1878. A. B. Meyer, Journ.
1874.
?
Casuarius
sp.
f.
v. pi.
all
;
15
A. B. Meyer, Sitzber. k. Ak. Wiss. Wien,
l)y
Rosenberg
;
but
Ixix. p.
it
is,
216.
of course, doubtfLd
whethei-
:
;
quite triangular in shape.
livid
casque low and
Face behind
Hind-neck
and fore-neck deep blue.
neck
(footnote)
Orn. pp. 200-203.
Adult. Bill short and pointed
it
417
apply strictly and only to this species, or to several, or whether tliey are nieaut for any
" Mswaar " at Andai, " Nhamdia " at Ilatam, " Wonggr " at Dorei.
form of Cassowary
make
id., Ibis, p.
pi. xix.
Native names: the I'olluwiug are given
they
248;
Scl., P. Z. S. p.
scarlet.
magenta-purple, posterior half
In the place where
in
'
the
There
is,
much
eye,
flattened
posteriorly, so as to
and head white.
Nape, throat,
Anterior half of naked lower sides of
scarlet.
Casuarius casuarius and
C.
however, nu certainty about
home.
its
umippendiculatus groups
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
140
the throat-wattles are attached there
Plumage black
colour as the rest of the fore-neck.
than in
those
previously described,
species
the
sometimes a small round knob of the same
is
10 inches long and very thick, forming
a sort of
;
feathers long and
over the
pendent
more
being some
tail
Legs brown,
train.
silky
12 to
and
bill
casque black.
Total length (according to Salvadori) about 1 m. 400 mm., tarsus 260 mm.,
gape 120 mm., inner claw 78
Young
mm.
Plumage brown,
(full-grown).
from
bill
colours on naked parts duller, the blue
all
being of a dull leaden shade, and the red replaced by orange-yellow.
Plumage dark chestnut-brown or bay-brown, each feather
Head and neck not bare.
strongly defined black bars.
(half-grown).
Yoiniq
crossed by several
Uniform daik brown, with two or three narrow longitudinal white bands on
Chick.
the body, and several similar lines on the neck.
This bird inhabits the Arfak Peninsula, and has been said to come also from
but this
is
.Tobi,
not supported by any evidence.
Eosenberg discovered
Guinea, where
mentioning
it
in
it,
is
this
called "
New
Cassowary near Andai, in the Arfak Peninsula of
Mswaar
" or
1871, Schlegel called
it
" Meswaar
'"
C. hennetti,
When
by the Papuans.
but stated that
it
differed
first
from
the latter in the coloration of the upperside of the neck, which, however, he did not
consider of any importance, and remarked that Rosenberg believed
it
to
be a new
species which he wished to call Casuarivs jMj^uonns.
Rosenberg
tells
us that the
first
glance at the type of his
shot by his hunter Aclimat (Achmed), convinced
Cassowaries of
New
papuaims, which was
that his former belief that the
Guinea were the same as those from Salwatti, the C. itnappendi-
was erroneous.
cidatus,
him
C.
Rosenberg further
during the breeding-season, which
consists of a depression
female was shot
states that the adult
from February to April, and that the nest
lasts
on the ground, lined with leaves.
He
had
all this
information,
however, from natives, and he repeats the old fable with which he also treated us when
describing the
Ceram Cassowary,
that
some eggs
(five,
he says) are placed in the
nest,
others (two) outside, so as to serve as food for the chicks, and that they are hatched in
Of
28 days.
The
had not a
knife.
little
trouble in keeping
it
first
off,
only wounded, and attacked the man,
and subsequently killed
])orei.
Bruijn sent skins to Europe said to be from Andai and Emberbaki, or Amber-
Beccari says
prints of a
it
it
near
occurs also on the mountains of Arfak, where he saw foot-
Cassowary at a considerable elevation.
The type
it
who
with his hunting-
it
Laglaize purchased one from Amberbaki, and A. B. Meyer obtained
baki.
that
the eggs he says that they are in every respect like those of C. casuarius.
adult bird shot by his hunter was at
of C. westermanni belongs certainly to C. j^apuanus.
was said by the missionary who sent
it
to
Sclater informs us
come from Munsinam (=Mansinam),
—
HON. W. KOTIISCHILD 0\ TlIK
near^Andai
doubt
Mansmam
and that
MUage
C.
to
of
where
also
th,s statement.
.s
rfEXr.s CASU.ARIUS.
Rosenberg obtained
Meyer
(Sitzber.
C. papnanns.
There is no reason to
Ak. Wiss. Wien, Ixix. p. 216) says that
k.
a sn.all island near Dorei,
where no Cassowary could Uve in a wild state,
Mansina,„ " is incorrect.
However, there is also a
Mansiman) a few miles south of Andai. and,
nK.reover,
tbei-efore the locality "
Mans.nam (or
westermanm does not
called
as
differ
from
„uun,s,
C.
suppose that the Mansinam, whence the
type of
Manaswan (which is, according to Meyer, the
alone bearing the
{of.
exist
the
141
map
name
of Mansinan.)-
It
it
C.
is
evidently quite unnecessary
^ocstermanni came,
proper name,
n.ore likely the
is
is
the island
principal
its
Mansiman of
village
the
Rosenberg and many others), which was quite
recently found
and was visited by William Doherty.
maps
ol
The eggs of
C. jiapiiunus in
still
to
Museum are rather smoother than most eggs
of the C. casnarius group and others, the granulations being rather
flat.
It is possible
that this flatness of the granulations is the rule in the eggs of the
C. papurarus group.
Those in the British Museum measure 134 86 and 130 84
the British
:
I
mm
:
Unfortunately, by carelessly leading Dr. Oustalet's description of
C.pajmraiNs cdwardsi,
erroneously gave Salwatty as a further habitat of this species,
and
it is
on the maps (Plates
15.
XL. & XLI.) which show
papuaxus
Casuakius
tlius
marked
the distribution of the genus.
edwardsi Oust.
Milne-Edwards's
Cassowary
(Plate
XXXV.)
1878. Casuarhis cdwardsi Oust., P. Z. S. p. 389,
xii.
p.
I.2.-.
;
Cab., Journ.
f.
Orn.
p.
203
pi. xxi.
(Dorey)
(editorial
note)
;
;
Salvad.,
Ann. Mus.
Civ.
A. B. Mever, Journ
f
G en.
Orn
p. 21)9.
1895. Casnarius jjo/mnnns (partim!) Salvad., Cat. B. Brit. Mus. xsvii.
Adult.
Plumage
black, feathers on
flattened posteriorly and black
;
bill
rump and
tail
much
long, pointed, and
p.
GOO.
elongated.
black.
Casque
16
much
Occiput, cheeks, and
below ear and cheeks a large black patch surrounded by
a broad
pink band, this pink band extends right round the back of the head below the white
occiput, but is concealed in the Iblds of the skin unless the bird is violently enra"-ed.
sides of face white;
Chin, throat, and fore-neck deep blue, in centre of fore-neck usually a round l)lack
Nape greyish black, rest of hind-neck scarlet.
wart with green centre.
Naked
lower sides of neck dull scarlet washed with
livid
pinkish purple, the spaces between
the folds black.
Reddish brown, feathers on fore part of body and back faintly
Occiput and sides of head bluish white, the occiput
barred transversely with black.
and
Face
part
of
whiter.
head round casque dark leaden grey, naked lower
being
Jiiv.
sides of
(half-grown).
neck pale flesh-colour
VOL. XV.
—PART
v.
No.
5.
;
legs yellowish olive.
December, 1900.
v
HON. W. EOTHSCHILD ON THE GENUS CASUAEIUS.
142
of head and face
Fore-neck and upper half of hind-neck dull
Occiput dirty brownish white.
black.
Crown
Casque not yet developed.
Juv. (three-fourths grown).
Large irregular patch between ear-hole and angle of gape buffy pink. Lower
Centre of naked lower sides of neck scarlet-mauve,
half of hind-neck rosy salniou-red.
Tlumage brown, curiously
bordered with a colour mixed orange and cherry-rose.
blue.
variegated with black.
Hab.
My
Low
living
country of
specimen performed as follows
inflating
:
neck horizontally, then contracts
It first extends its
same time
N.W. Dutch New Guinea.
it,
and then makes a
feathers of the back and hind-quarters out and
it,
like other
the
at
it,
short and deep
Cassowaries, spreads the
snarls viciously.
described above
three-quarter grown bird
The yonng
least twelve
series of at
Sometimes, when very angry,
grunting notes.
short and bends
it
is
not
yet
old
enough
to
perform these antics.
Milne-Edwards's Cassowary was
Oustalet
described by
first
from a male
The
near Dorei, and brought to Paris by Messrs. Eaffray and Maindron.
noticed
by
coloration
M. Oustalet
;
form
while the form of the helmet
is
the
of
The
and neck.
naked parts of the head
of the
very conspicuous
the
in
principally
are
more
helmet
latter
killed
differences
and
indeed,
are,
likely to be an
the
individual
character.
I
cannot believe that C. edwardsi
subspecies of the latter, then
"identical" with
papuanus; but
it
is
a
the two forms must iidiabit different areas, and
it
is
is
C.
very likely that one of them frequents the lowlands, while the other lives
if
among
the
hills.
An
flat,
egg laid by a living bird in
paimanus
like those of C.
my
possession has the network of granulations rather
abnormally small, measuring only 125
16. Casuarius LORiiE Rothsch.
1895. Casuarius
j}icticollis
1898. Casuarius
lorice
:
8G
It
(Plate
Salvad. (nee Sdater), Cat. Birds Erit.
viii.
p.
p.
is
rather pointed and seems
mm.
Loria's Cassowary.
Rothsch., Novit. Zool.
1899. Id., Bull. B. O. C.
Museum.
in the British
XXXVIII.)
600 (description
Mus.
xxvii. p.
S. p.
775 (June 20tb
!).
513.
hi (Jime 21st)
;
id.,
P.
Z.
;
appeared
Oct. 1st).
Adult. Bill short and pointed, castiue low and
and mesial
line of
fore-neck scarlet.
blue.
neck
much
flattened
Throat
posteriorly.
neck dull pink, with three narrow longitudinal lines of blue,
In front of the ear a small patch of
Head and hind-neck
bright
blue.
Legs horny grey.
rest of
magenta, above the ear
Anterior portion of naked lower sides of
livid purple, posterior portion scarlet, the
black, long and silky.
livid
whole surrounded with blue.
Iris hazel.
Plumage
HON. W. EOTHSCllILD ON THE GENUS CASUAKIUS.
The
the
from gape measures
bill
Museum
of the fullv adult male in the
total lenoth
according to Salvadori.
Weiske) measures :—
11^0
Genoa
is
about
H
m.,
mm., the tarsus 244 mm., claw of inner toe 117 mm.,
iMy adult
liill
of
143
Upper Brown River
the
bird froui
(collected by
from gape 122 mm., tarsus 246 mm., claw of inner toe
mm.
70
Young (haU-grown) Ivufous-brown, each
feather
.
marked with narrow
transverse bars
of brownish black.
Hah. Highlands of S.E. New Guinea.
Casuarius lurin- was found by Signer
Moroka in the highlamls of the
Owen Stanley Mountains in Britisli New Guinea. The fully adult specimen obtained
Mr. Loria made a coloured sketch of
by Loria is now in tlie Museo Civico of Genoa.
I.oria at
me
the head and neck, wliich was kindly lent
is
by Dr. Gestro, and from which our plate
taken.
It is this
same bird which
is
'
Catalogue of Birds,'
which shows the red of
bird, a skin
Another adult
described in the
tlie
vol. xxvii. p.
neck well, was sent
Mr. Weiske from the mountains on the Upper Brown River.
I
601.
me by
to
cannot possibly con-
found these red-necked individuals with Cnsnnrius picticolUs, which has the neck blue
with a small red spot in the centre of the fore-neck, and a red chin when young.
C. lorias
to
(/.
evidently replaces C. picficoUis in the highlands, the latter being, according
our present knowledge, only found in the coast-region.
T"he second example mentioned in the list of specimens in the
c.)
Casu.\kius picTicOLLis Scl.
1871. Casuarius
iJtcticnU'is Scl.,
1875. Scl, P. Z. S.
1877. Gould, B.
1881. Salviid.,
New
Mem.
1882. Salvad., Aim.
1893.
1894.
(?)
(?)
p.
P.iinted-necked Cassowary.
Rep. Brit. Assoc.
Guinea,
Pv.
v. pi. 14.;
I^lus.
Civ. Gen.
xviii. p.
Sehalow, Jonni.
f.
Orn.
p.
1881.
Casuaiins kaiipt
[wm
Aihdt. Plumage black
25 (egg)
black.
p.
p.
;
319 (puUus, Miliie Bay)
ibid. p.
213, pi.
3, p. 29, pi.
fig.
ii.
9 (head)
ii.
H-s.
1,
Ibis, p.
p.
499;
;
5(j;)..
sides of
;
bill
pointed.
Chin, throat, fore-neck, and
In centre of fore-neck a deep red patch.
Lower hind-
neck bright rose-purple.
Total length about 1 m. 400 mm., tarsus 245 mm,,
mm.
2 (egg)
iii.
;
Occiput and auricular region pale greenish blue.
inner claw 125
;
casque high and depressed posteriorly, black
nape of neck deep bright blue.
neck pale blue. Lower naked
;
;
600.
Rosenberg!) Sharpe,
;
XXXVI.)
415, no. S; Salvad., Orn. Pap. e Mol.
A. B. Meyer, Abh. Miis. Dresd. no.
1895. Salvad., Cat. B. Brit. Mus. xxvii.
(Plate
Forbes, P. Z. S. pp. 307, 315, 31(3 (anat.)
Ac. So. Tor. (2) xxxiv.
'
138.
\).
81, pi. xviii. (Discovery Bay)
Catalogue of Birds
under that name.
1 could not find in the collection, at least not
17.
'
bill
from gape about 120 ram.,
HON. W. ROTHSCHILD ON THE GENUS CASUAEIUS.
144
different
the two most
:
patch in centre of fore-neck;
red, nearly joining the red
Chin and throat dull
(1)
bird varies considerably, and I describe
The immature
Immature.
rest of naked parts as in adult, only duller in colour
plumage black, completely
;
mingled with brown feathers.
feathers.
Chin, throat, fore- and
hind-neck, Hnd occiput dull blue, base of hind-neck orange.
In centre of fore-neck a
Plumage brown,
(2)
sliglitly
mixed with black
large horseshoe-shaped red patcli, lower naked sides of neck dull purple.
Juv. (one-third grown). Uniform pale brown.
New
Hah. Coast-region of Briiish
In 1875 (P. Z.
"In
C.
Guinea.
85) Dr. Sclater characterized
S. p.
ivestermanni
the throat
(pi. xix.)
the middle of the throat
red,
is
bird as follows:
—
blue and the liinder part of the neck
is
In the new species, which
deep orange-red.
tliis
1
propose to
call C.
picticoUis (pi.
and the hinder part of the neck bright blue.
xviii.),
There
are, besides, minor differences, which will be evident on comparing the two drawings.
Now,
so
constant; and
I
The
species."
naked parts of the Cassowaries are quite
can hardly doubt therefore that we have here to deal with different
know, these colours
as I
fixr
original
specimen came from Discovery Bay.
lived for about twenty-nine
in
description of
i)iGticollis
C.
that of a mountain form
mouths
in
in tlie
which
Catalogue of Birds
•
It died,
and
from
in
is
after
now
Milne
having
preserved
Bay.
Museum
the British
The
'
is
consider to be specifically distinct, and which
I
have separated as Camaniis hriw, as
of
the Society's Gardens,
Other individuals are known
Museum.
the British
in the
had been able
I
to
I
examine several specimens
it.
18.
Casu.\rius picticollis HECKi Rothsch.
Heck's Cassowary.
1899. Casiiurius picttcoUis hecki Rotlisch. Bull. B. O. C.
id.,
Adult.
P. Z. S. p. 773 (June
]>ill
short and pointed
crimson, bordered
This form
chicks
differs
casque low and flattened
with indigo-blue.
Legs brown.
Young and
;
;
Bill
l~tli), p. Ivi
ijosteriorly.
Naked lower
(June 21st)
;
Occiput Nile-
sides of
neck bright
Cheek-wattles large and balloon-shai)ed when
and casque black.
unknown.
from
C. picticollis in
the absence of red on the throat and chin, in
the much darker blue of the head and neck, and
purplish-red naked lower sides of neck.
Uah. German
(May
XXXVII.)
ajjpearei Oct. 1st).
and neck deep indigo-blue.
blue, rest of head
extended.
.'20th
viii. p. xli.\
(Plate
New
Guinea.
in
the bright crimson instead of
—
;
HON. W. KOTHSCHILD ON THE GENUS CASUARIUS.
Nothing
kuowu
is
My
of Berlin.
of this form except one living individual in the Zoological Garden
original preliminary description of
" This bird bi'ars tiie
does to
C.
paimanus.
A
greenish blue.
dark crimson.
form
14G
same relationship
it (/. c.) is
as follows
to C. ijicticollis that C.
:
papuanus edwardai
Tlie throat and liind-neck are deep indigo-blue.
small round bhu k wattle on the fore-neck.
Casque and plumage similar
Occiput pule
Lower
neck
sides of
to those of C. picticollis."
m
named
honour of Dr. Heck, Director of the Zoological Garden of
Berlin, who kindly permitted Mr. Keulemans to make a drawing of the bird.
Tliis
19.
is
The Mooruk.
Casu.arils be.nnetti Gould.
1857. Casuarius beuneiti Gould, P. Z. S.
& Mag.
1858. Gould, Ann.
1858. J. E. Gray, P. Z. S.
1859. Bennett, P. Z. S.
Nat. Hist.
205, Aves,
p.
Naturalist in Aiistralas.
p.
1862. Scl., Trans. Zool. Soc. Lond.
18G3.
p.
299
p.
pi.
pi. iv.
ix. p.
359,
Nat. (Micrb.
18G4. Scl., P. Z. S.
234 (incubation),
ix. p.
p.
323
Scl.,
P. Z. S.
ii.
vii. p.
1871. G. R. Gray, Hand-list,
p.
iii.
5G1
p.
Garrod, P. Z.
pi. Ixii.
518
S. pp.
1876. Ramsay, P. Z. S.
p. 2,
p.
Schleg., Jaarb. zool. Genootscli. Nat. .\it.
;
1880. Layard, Ibis,
]).
303; Powell,
Mem.
R. Ac. Se. Tor.
1882. Salvad., Ann. Mus. Civ. Geu.
p.
no.
p.
4; Beuu.
&
Sclat., P. Z. S. p.
p.
112;
519
O.
20 (egg)
p.
Mus.
;
P. Z. S. p.
(2)
493;
xxxiv. p. 214, pi.
xviii. p.
415, uo. 9;
ii.
id.,
fig.
10 (head)
;
Orn. Pap. e Mol.
p.
11
iii.
p.
" iu
501
iii.
;
p.
438;
;
:
Paris, p. 256.
1869. Cassowary from the Solomon Islands (error!), Hutton, Ibis,
Moornp
Ac. Soc.
(corr. of locality)
84 (period of incubat.); Sliarpe, Cat. Ostcol. Spec. R. C. Surg.
1896. Oust., Nuuv. Arch.
black.
Mem.
9850;
1892. Ost, Zeitschr. Ool. pp. 15, 91; Schalow, ibid.
f.
Jouan,
122;
5;
posteriorly,
Gatii.
;
470, 614 (anatomy)
p.
Plumage
;
178;
1879. Scl., P. Z. S.
''
235
210; Bennett,
p.
(descr. oF pullus), pi. xlii.;
1875. Scl., P. Z. S. pp. 85, 87; Scl., Rep. Brit. Assoc,
:
pp. 212,
;
1873. Schleg., Mus. Pays-Bas, Strid/iiones,
Native name
Ibi.s,
;
271 (successfully reared)
18GG. Seldcg., Zool. Gart.
1894. Id., Jonrn.
72, 73; Sel.,
;
1865. Gould, Handb. B, Austr.
1891. Evans, Ibis,
pis.
198
Scl., P. Z. S. p.
1881. Salvad.,
;
;
1G2 (egg);
213,
XXXIX.)
;
144 (egg)
pi.
129
pi.
32; GouUl, B. Austr., Suppl.
p.
18G0. Bartictt, P. Z. S.
Mag.
i.
(3)
271, Aves,
p.
269, Aves,
p.
(Plate
New
Bill long,
Britain (corrupted:
p.
352.
"Mooruk").
arched and pointed.
Casque high, sharply depressed
the various ridges being sharply defined.
Back of head, occiput, and
;
HON. W. EOTHSCHILD ON THE GENUS CASUARIUS.
146
upper hind-neck black-blue
;
lower hind-neck dark blue.
Chin, throat, and sides of
Below the ear-hole and
fore-neck deep bright blue, rest of fore-neck blackish blue.
reaching to the gape
naked lower
is
sides of
Upper half
a large irregular patch of very pale blue.
of the
neck black, with purplish-pink streaks between the wrinkles,
lower half purplish pink with black spots.
Total length about
1
m. 330 ram.,
from gape 110 mm., tarsus
bill
-ilO
mm., inner
claw about 73 to 90 ram.
Head and hind-neck
Chick.
and
thioat
yellowish
buff.
bright
Fore-neck and
with black.
Rest of body crearay
with
running along
rufous
mingled
rufous
buff.
Chin
spots.
mingled
buff
longitudinal black bands variegated
back, and two
tlie
black
neck rufous
of
sides
Five
with
bands down the flanks and
black
thighs.
Hah.
New
The adult
but
it is
Britain.
bird here described was sent
impossible to say
if
me
is
New
an inhabitant of the island of
and unscientifically
officially
plumage
the bird was adult then or not, for though the
was black, the casque was undeveloped, and the colours were
The Mooruk
Webster;
four years ago by Captain Cayley
dull.
Britain (now most unreasonably
renamed " Neu Pommern
was
It
").
for the first time
made known in a letter from Mr. George Bennett, communicated to this Society
who proposed the name Casuarius hennetti for the new bird, and reproduced
a drawing made by Mr. G. F. Angas from the live bird, which was not quite mature.
The letter reads as follows
" My dear Gould, I send you an account of a new species of Cassowary recently
by Gould,
:
brought
Sydney by Captain Deolin in the cutter
to
'
Oberon.'
It
was procured from
the natives of
New
of
tlie
3 feet to the top of the back, and 5 feet
is
rufous mixed with black on the back and hinder portions of the body, and raven-
bird
is
Britain,
where
it is
black about the neck and breast
:
known by
the
name of Moorirk.' The height
when standing erect its colour
'
;
the loose wavy skin of the neck
is
beautifully
coloured with iridescent tints of bluish-purple, pink, and an occasional shade of green,
quite different from the red and purple caruncles of the Casuarius galeatus
and
legs,
;
the feet
which are very large and strong, are of a pale ash-colour, and exhibit a
in the extreme length of the claw of the inner toe on each
remarkable peculiarity
foot, it
being nearly three times the length which
other toes.
a
This bird also
helmet-like
protuberance
character of and
of the
bill
differs
from
it
C. (jaleatus in
obtains
the claws of the
having a horny plate instead of
on the top of the head, which callous plate has
resembles mother-of-pearl darkened with
differs
in
considerably from
that of the
Emu
black-lead.
the
The form
{Dronueus novce-hollandice),
being narrower, longer, and more curved, and in ha\ing a black and leathery cere
at
the
base,
and behind the plate
of the
head a small
tuft
of black hair-like
—
HOX. W. ROTHSCHILD ON THE GE^a^S CASUARIUS.
feathers,
which are continued
in greater or lesser
147
abundance over most parts of the
neck."
The
female by laying eggs.
;
and
this
be often kept as a pet by the natives of
and
its
neck seems
There
is
to
Dr. Bennett, and proved itself to be a
most Mooraks, much more gentle and tame tlian
seems to be a peculiarity of the Mooruk, which is said
It was, like
most other Cassowaries
to
London by
original specimen was sent to
New
Britain.
bears itself less uju-ight.
It
be shorter than that of other Cassowaries.
a very long and amusing account of
its
habits in captivity in Dr. Bennett's
Gatherings of a Naturalist in Australasia,' accom])auied by a figure by Wolf.
In 18(33 the Mooruk hatched a nestling in tlie Society's Gardens.
The female
began to lay in the middle of March, and laid half a dozen eggs at intervals of about a
'
The incubation, according to Dr. Sclater,
The young did not live nnn-e than a
52 days.
week.
time described (and figured, P. Z.
S.
olive tul)ercles, but the eggs laid in
They do not
differ perceptibly
day.
were
Tlie eggs
for the
lirst
1858) as being of a pale olive colour with darker
London
apple-green colour (see figure, P. Z.
lasted once 49 days, the next time
when
sh(jw that they are
fresh of a bright
18G0) with darker green glazed granulations.
S.
from the eggs of Casuarius casuarius and
allies,
though
the glazed caruncles are somewhat less continuous and not so thickly and equally
The following measurements have been
131:90 to S3, 150:88, 135:89, 13(5:90,
distributed over the entire surface of the egg.
obtained:— 137: 88, 128:81, 141 to
152 87 mnr. The lull fresh-laid egg weighs about 22 ounces.
:
that there
hardly any difference between the eggs of
is
Schalow's desciiption of the colour
The smooth ones
faded ones.
is
('.
not from fresh eggs
(see P. Z. 8.
It will
thus be seen
and
C. cusibarins.
heiuietti
—
all
he saw were evidently
1858 and 18G0) are
first-laid
eggs, the
roughly granulated ones the properly developed eggs.
The
notes of
my
live
specimen
— evidently
a male
— are
usually a low and short
piping note, reminding one of that of a large chicken, and when a
this note is louder
and
short,
utters
and ino;e
first
some
phriiitive.
higher, tlien
When much
excited
it
makes
little
its
excited
neck quite
some lower barking sounds followed bv some
snarling ones.
'J'he
eggs of Casiuiriiis bennetli
hardly two being alike.
that
due
is
One
Cassowaries" eggs.
one
is
almost
like the
131
:
90
smooth
eggs of
137:90 mm.
to
83
They
exposure,
to
C.
as
in the British
Museum have
are of different shades of brown, but, needless to say,
we know the
fresh
eggs
are
green like
all
other
has very few, large and highly raised glazed granulations,
(like
first-laid
pcquutnus.
The\
eggs of Cassowaries
measure:
generally), one
exactly
141:89, 127:80, 140-5:90, and
Schalow gives 137:88 and 12S'5:8L5 mm.
mm.
a very different surface,
Ramsay
gives 141
to
HON. W. EOTHSCHILD ON THE GENUS CASUAEIUS.
148
20. Casuarils bennetti maculatus, subsp. nov.
Casque high, much depressed posteriorly, and sharply ridged.
bill
long, arched,
specimen of
as a second
blue
;
;
inner claw long.
Naked lower
extending to the gape
is
first identified
On
and the fore-neck paler and brighter blue.
H
a large round patch,
is
(This bird was
of neck
me
the lower
inches in diameter and of a rosy-purple
rosy
purple
a large white patch
washed
sides
by
Occiput and hind-neck bright indigo-
C. picticolUs liecki.)
sides of head, face,
part of fore-neck
colour.
and pointed
Plumage black
:
from
and
below the ear-hole
with rose-colour.
Hah. unknown.
may eventually prove
we do not know the
This form
so long as
Cassowary, and the
and describe
is
be only a colour-aberration of
exact
C. hennetti
habitat of every species and
full extent of their individual variation, I consider
this bird as a subspecies of C. hennetti.
Society's Gardens,
and
to
and was
not so wild as
my
fully adult
when
specimens of true
it
The
was imported.
type
It
is
it
now
;
but,
subspecies of
right to
name
living in the
seems to be a male,
C. hennetti usually were.
—
149
[
Part
]
Phylogeny of the Palteognathse (Ratitse and Crypturi)
and Neognathse (Carinatse). By W. P. Ptcraft.
U.— On the Morphology and
Table of Contents.
Page
Pferyhsis
Of Adult
„
Embryo and Nestling
Structure of Feathers
The
of Casuarius
.
Teleoptiles
Summary
151
Brain
247
159
Eye
247
162
Jacobsou's cartilages
247
162
Neossoptiles
Adult and Nestling Kemiges
of Pterylological Characters
....
Page
Nervous System and Sensory Oryans.
Alimentary Canal and Appendages.
165
Buccal cavity
248
166
Intestinal convolutions
248
168
Liver and Gall-bladder
251
Respiratory Organs.
Lungs
252
171
Air-sacs
252
195
Windpipe
254
Choanaj
256
Osteology.
Skull of Adult
Nestling
Some
points of significance in the Strathious
206
Palate
Circulatory System.
256
Vertebral Column
20S
Heart
Kibs
215
Carotids
Sternum and Pectoral Arch
216
Pelvic
Pectoral
Pelvic
225
Arch
233
Limb
235
Limb
257
Bursa
Cloaca,
Fabricii,
and
-38
Skin-muscles
Muscles of
Wing and
Shoulder-girdle
.... 238
242
Thigh- and Leg-muscles
-
genital
System.
257
Cloaca
Bursa
258
Copulatory Organ
259
259
Development
Myology.
Urino
Phylogeny of
the Palaogiiatha
Keys
to
2-59
266
Summary
Genera and Species
267
Introductory.
present investigation was undertaken at the request and with the co-operation
furnished the bulk of the
of the Hon. Walter Rothschild, M.P., who has furthermore
The collection of
work.
material, and in numerous other ways greatly aided my
The
skeletons in the British
Museum
of Natural History
has proved a source of great
am besides indebted to Mr. Beddard, Dr. Gadow, Dr. Forsyth Major, and
possession or charge, and I take this
Prof. Howes for the loan of specimens in their
opportunity of recording my thanks.
wealth.
1
so possible, the relaobject of this investigation has been to ascertain, as far
"
" Struthious forms, and the position of these
tions of the CasuariidcB to the remaining
z
December, 1900.
PART V. No. 6.
yoL. XV.
The
—
ME. W.
150
PYCEAFT ON THE MORPHOLOGY AND
P.
with regard to the " Carinatce."
That
this
attempt has not been
fruitless will
probably
be admitted.
From
the nature of the aim of this paper,
of investigation adopted
Moreover,
will I trust be agreed
about the best possible.
is
characters which bear directly or indirectly
group.
it
The
that the
been to
desire has
at
tlie
serve the ends 1
upon the problem of the phylogeny of the
end of
had
work, and from which
this
Hence
view.
in
in
the memoirs
much has been
extracted to
may be
The memoirs of Garrod, Gadow,
the majority of the omissions which
noted in these pages are not accidental but of design.
Fiirbringer, Meckel, Cuvier, D'Alton, Parker, Beddard, Lydekker,
milestones which
this group.
mark
and
There seemed
to
me
to
be no reason or end to be gained
add a few courses more
The
to
to the structures
rejection of the old terms Ratitce
be warranted.
The
and
to
endeavour
which they have reared.
and
Ratite condition
in transcribing
appeared more expedient
it
to set myself the task of recording the substance of their achievements,
me
others are the
the progress of the knowledge which has been accumulated on
the information which these individually impart, but rather
to
select those
have dealt in detail upon the anatomy of the group would have
to
been to repeat a vast body of facts already well known and ably treated
enumerated
scheme
is
Carinatce,
which I now propose, seems
admittedly a secondary one, and the
some " Carinatm " are also " Ratitce."
terms now suggested are mutually exclusive, and based upon a primitive character.
is
objectionable, in that Hesjjerornis and
Furthermore, as will be shown, the change
this account they are to be preferred.
to
be attributed to a desire to
effect a
suggested but for the need, and
the group hitherto
known
change for change sake:
seems
it
perhaps have been so very contradictory,
there
is
to
as " Eatitce."
me
it
to
name
The
On
is
not
would not have been
a real need, to include the Crypturi with
To have
for, since
retained the old terms would not
some " Carinatce " are
no reason why some Ratitce should not be " Carinatce
A
"
!
really "
Ratitm"
further discussion
of the subject will be found in the summary.
In conclusion, I wish to record
work
entrusted this
to
my
my
grateful thanks to Mr.
Rothschild for having
hands.
PTERYLOSIS.
The
description of the pterylosis of the Palceognathce
by an enumeration of the apteria.
nevertheless
of considerable
Though
is
most
easily
accomplished
these are but few in number, they are
importance, inasmuch
as
they
disprove the
prevalent
notion that the feathers in the '•'Ratitce" are evenly distributed over the body.
has, however, long been
e. g.
Struthio, Rhea,
The
known
It
that apteria occurred in embryos of certain forms,
and Apteryx.
descriptions of the Oil-gland, Rhamphotheca, Podotheca, and of the structure of
the feathers, both macro- and microscopical, will be found in this section.
: :
PHYLOGENY OF THE PAL^OGNATH^ AND NEOGNATH^.
Pterylosis
a.
of
151
the Ad^ilt}
Casuariid^.
Casuarius casdarius Linn.
Ajjferia
A. spinale.
—
—This was wanting.
—This includes the
A. mesogastrcei.
and extends backwards
characteristic and
well-marked sternal
callosity,
within a short distance of the cloacal aperture.
to
It is
widest on the abdomen.
A. trunci laterale.
— Represented
small truncal area at
The head and more
its
by the naked under surface of the wing and a
base.
of the
or less
neck
the adults of this
are, in
The extent and nature
species of the genus, bare, and brilliantly coloured.
coloration have already been dealt with by the
and other
Hon. Walter Rothschild
of this
in the
first
part of this Memoir.
Pteryke
P. alaris (PI,
—
XLV.
10).
fig.
—
It is possible to distinguish in
The former
coverts as in Neognatha'.
elongated porcupine-like
as follows:
— mid-digital
quills.
(1)
are represented
These are
six
in
this
both remiges and
by the characteristic
number and
distributed
(2) representing the primaries,
and metacarpals
and
cubitals or secondaries (3).
The
cubital series, at
in Neognathce,
the ulna than
appears not to extend backwards to the elbow as
not farther
of the spine-like remiges being attached
the last
its
first sight,
proximal fourth.
Beyond
this,
however, are three moderately long
feathers occupying the position of remiges, but bearing each an aftershaft.
at the distal
end of the
series,
the position
of a remex
—
an
aftershaft.
down
Similarly
have, in front of the mid-digital, a feather occupying
we
possibly representing the addigital
—but
small and bearing
This, like the three cubitals, appears to have a major covert, as also
have the other spine-like remiges.
are of considerable length.
The
Tiie mid-digital
is
very small
;
the other remiges
longest of these are the 1st metacarpal and the
1st
cubital.
The remiges are remarkable in that they are open at the free end (see page 166).
They are cylindrical, of irregular thickness and more or less crooked.
The tectrices are not distinguishable from the rest of the body-feathering. All bear
aftershafts.
'
As
" adult "
Coverts representing the tectrices majores of the JVeognathce are certainly
we have
included nearlj- full-grown as well as full-grown specimens,
yet attained the characteristic adult livery.
i.
e.
those which have not
152
ME. W.
.
present,
and bear the same
and marginales
There
is
PYCRAPT ON THE MOEPHOLOGY AND
P.
relation to the remiges.
Other coverts
t.
medioe, miiiores,
—are not distinguishable.
no ala spuria.
Neither in
this,
nor in any other species of the Palceogiiathce are there any semi-
plumse, plumulEe, or
filo
plumulse.
The absence
There are no
than of the two former.
of these last
The
rectrices.
is
oil-gland
is
of more significance
wanting.
Eyelashes are present and well developed.
The Rham2}hotheca
of the upper
jaw
rhino- and small lateral gnathotheca.
theca, but
slit-like,
The
is
The
nasal fossa
is
The
covered only by the integument.
is
Podotlieca
(fig.
2
not protected by the rhampho-
anterior nares are impervious,
155) has the acrotarsium clothed, below with broad scutes,
a, p.
;
the acropodium with scutes.
greatly elongated, to serve as a
The wing
C.
compound, being composed of a small median
slope obliquely forward, and placed at the extreme end of the fossa.
above with large granular scales
toe
is
is
weapon of
The claw
of the inner
offence.
not armed with the blunt claw found in
many
other species.
casuarius heccarii (Sclater).
The
pterylosis of this subspecies agrees with
that of jthe typical form C. casuarius.
In one specimen I found 6 remiges, the additional quill representing a primary.
C. casuarius sclateri.
The
pterylosis of this agreed with tlie
two foregoing.
In one half-grown specimen
the apterium spinale could be faintly traced from the inner surface of the skin.
The
wing bore a long blunt claw.
C. casuarius salvadorii.
Apter. s^nnale.
—Though
not of great extent, in this species was well defined.
It
extended cephalad as far as a point corresponding with the exit of a vertical line
passing through the body from the posterior end of the sternal callosity, and caudad to
the middle of the pelvis above the acetabulum.
its
Its total
length was about 8 inches,
width about \ inch.
A. mesogastrcei.
— Very
narrow, and bounded by degenerate [feathers, indicating a
tendency to increase the width of this space, and recalling similar conditions in the
Carmat(B.
It could
be traced backwards to within a short distance of the cloacal
aperture.
A. trunci later ale.
—
— Represented by the naked under
Pteryla alaris.
As in C. casuarius.
The wing was armed with a claw.
surface of the wing.
a
::
PHYLOGENY OF THE PAL^OGNATH^ AND NEOGNATH^..
153
C. CASUARIUS VIOLICOLLIS.
C. UNAPPBNDICULATUS,
A. spinale.
— As
A. mesogastrcei.
in C. casuarius sahadorii.
—This
was much more
clearly defined than in
C.
casuarius.
Its
width exceeded that of the apterium spinale.
I
have not been able
sequently
am
examine spirit-specimens of nestling Cassowaries.
to
Con-
unable to say whether there
is an oil-gland, as in Drommis (p. 154)
whether the general pattern of t!ie body-coloration is continued on the podotheca,
and the integument covering the nasal fossa features which obtain in Droma'us.
I
;
or
:
One conspicuous
difference
between the nestling and the adult Casuarius
that the head and neck are densely feathered in the former.
by a horny plate covering the roof of the
DEOMiEUS Nov^-HOLLANDi^.
Ajiteria
The casque
is
indicated
Casuarius, but
is
relatively
skull.
(Nestling.)
—This wanting.
A. mesogastrm. — In form and
is
relations this agrees with
smaller.
A. trimci laterale.
Pterylce
P. alaris.
the fact
—
A. spinale.
much
is
— As
in Casuarius.
—
— There are
17 remiges in
These remiges
digitals.
in
Of
all.
the adult
these,
are
10 are cubitalsand? metacarpo-
peculiar,
lacking the calamus and
possessing an aftershaft.
The
tectrices are
arranged in obliquely transverse rows, as in Bhea
are not sharply separated one from another as in that genus.
(p.
156), but they
Furthermore, they are
not separable into major, median, minor, and marginal series as in Neognathw.
extend outwards on to the manus.
carpal series.
Digit II.
That
free.
is
They
All the primaries apparently belong to the metais
to
say,
so
much
of this vestigial digit as
remains projects freely beyond the feathers, and bears a claw as in ArchcBOpteryx,
In these last the claw appears in the embryo.
Opisfhocomus, and the Gallina'.
There
is
no ala spuria
Bhampliotheca.
caudad,
is
The
and rectriccs are also wanting.j
—That of the upper jaw resembles that of Casuarius.
sharply distinguished from a thin
impervious,
beak.
;
They
are
placed.
The
position
cere
of the
in
which the
nostrils
is
The rhinotheca.
nostrils,
which are
about the middle of the
are not protected by an operculum.
sheatli of the lower
median and two
lateral.
jaw
is
also
The tomium
composed, as in Casuarius, of three pieces
—
bears slight traces of denticulations similar to
154
ME. W.
P.
PTCEAFT ON THE MOEPHOLOGT AND
those described by myself in the
Tinamous [8i]
i.
The young
of Casuarius bear traces
of similar denticles.
Podotheca.
—
^The acropodium
is
made up
of small reticulate plates which gradually
fuse to form ten large transverse scutes, covering
middle of the
is
The border
series.
clothed with
these extend
;
proximal half of both outer and inner toes
elongated as in Casuarius.
It is interesting to
—the
downwards
The planta
so as to clothe the
remaining region of these, like the
The inner claw
being protected by large scutes.
toe,
in the
region, between the acrotarsiura and the acropodium,
tiny granulate plates
whole of the middle
These are widest
its distal third.
is
not
covered with small reticulate plates.
is
note that the podotheca and the cere are mottled with black
pigment, so as to carry out the general scheme of coloration which characterizes the
This mottling is most distinctly marked in a ripe embryo
feathers.
than
earlier
and in the nestling, they are not nearly so
this,
The Uropygmm is distinctly
much more easily demonstrable.
embedded
partly
in the skin
traceable in this nestling
Here
it is
(fig.
2
In a stage
b).
distinct.
:
in the ripe
embryo
it is
represented by a small rounded oval mass
and lying immediately above the cloacal aperture, as in
Apferyx.
Neither in this nestling nor in the embryos
egg-tooth, as
is
usually borne
found in the Neognathce
Perhaps we should be correct
colour.
becoming absorbed
;
there any trace of a separate, detachable,
is
produced into a small conical elevation of a whitish
in the embryos,
is,
is
but the region of the beak in which this
after hatching.
in regarding it as
In the Neognathce
a non-detachable egg-tooth
it
detachable, and falls off
is
after this event,
E.H E
I
D^.
Rhea ameeicana.
Apteria
A. spinale.
portions
—
—In one adult and
— more or
less distinctly
a three-months old nestling this was divided into two
— an anterior and a posterior
of this tract in some Tinamous,
cordiform in shape, with
2
in., its
in. in
tail.
1 in. for
It
base towards the head.
the rest of
other
A
Its
more or
;
thus recalling the form
The
anterior space was
greatest width was nearly
less perfectly
feathered area
length divided this from the posterior apterion which
was at
anterior aptei'iou
J'n
Calodromas elegans.
greatest length about the same.
of about 4
to the
its
e. g.
is
continued
about f in. wide, narrowed to 1^ in., contracting again to
course.
In another adult and three-months old nestling the
first
its
was very small.
and seven young
adults,
birds
ranging from embryos
specimens, the anterior space was wanting.
^
Figures in brackets refer to Bibliography at end of paper.
to
half-grown
PHYLOGENT OF THE PAL.EOGNATH^ AND NEOGNATH^.
155
Fig. 2.
A.
B.
D.
E.
a.
— Left foot of a nestling Casuariiis casvaruts schteri,
— Left
anterior aspect,
Casuarius
for
C.
— Left
A.
to
foot,
of
lorice,
show the arrangement of the
scutes of the podotheca.
comparison with A, to show the large hexagonal
plates clothing the proximal portion of tho tarso-metatarsus.
foot of a
newly hatched Dromaiis novce-lioUandic,
to
show the arrangement
podotheca, and the curious continuation of the pattern of coloration of the hody
— Left
D.
foot of a ripe
of feathers (/)
E.
— Left
foot of
of the scutes of the
down
to the toes.,
emhryo of Rhea americana, to show the arrangement of the scutes. Note the presence
— temporary in this species, permanent in if. darwini.
on the tarso-metatarsus-
an embryo Ajyieryx australis mantelU,
to
show the form
of the scutes of the podotheca.
ME. W.
156
A. trunci
backwards
width
laterale.
—This
PYCRAFT ON THE MORPHOLOGY AND
occupies the whole under surface of the wing, extending
corresponding to the level of the free end of the scapula.
to a point
It extends dorsad,
considerable.
is
P.
above the scapula, about
I in.,
Its
and ventrad,
and upper end of the thigh are
small and sparse, suggesting the formation of an apterion, thus tending to form a
distinct 2^teri/la femoralis as in the Neognathw.
A. mesogastrcei. This includes the sternal callosity, vvhich is 3| in. long and
below
it,
about 4
The
in.
feathers over the front
—
Immediately caudad of this callosity the space narrows to 1| in., and
2 in. wide.
again to about i in., which width is sustained for the rest of its length till it ceases
about 3
in. in
front of the cloacal aperture.
Pterylce
P. cavitis.
—
—Tlie
region of the eye
is
surrounded by a bare space
;
so also is the
The rim of the aural orifice is surrounded by feathers
the aperture. The throat is bare along each side of the mandible.
external aperture of the ear.
which serve to close
There are well-developed eyelashes.
Pt. alan's.
—The
wing
Metacarpo-digitals 12
=
The arrangement of the
is
eutaxial and of great size.
metacarpals 7
;
addigital 1
and coverts
quills
in
;
Remiges
28.
mid-digitals 2
;
Cubitals 16.
pre-digitals 2.
Rhea appears
the wing of the
to
be
unique, at least in so far as the arrangement of the cubital remiges and the coverts
are concerned.
The
The
pollex only bears a claw.
cubital remiges are spaced about as wide apart as in the
of about the same relative
which they
The
seem
1st
to
diflFer
size, e. g.
considerably
rows
:
and 6th almost reach
it,
have been carried away
Stork or Eagle.
1 to 6
do not
There
rest,
as
wing of a Neognathine
one point, however, in
is
the others, on the ulna.
the 3rd and 4th are most distant from
by the tension of the ala
membrana
it.
They
posterior, stretched
in the angle between the manus and forearm.
The
cubital remiges are rather shorter than their major coverts
are longer.
:
those of the hand
This difference in length between the remiges and their major coverts
probably an indication of degeneracy.
fact that in the
young chick the
first
is
It is interesting to note, in this connection, the
cubital fiight-feathers are not remiges, but major
coverts.
The metacarpo-digital remiges are arranged much as in the Weognathce, but
somewhat farther apart, so that they tend to form pairs with their major coverts.
Like the cubital remiges, they are of great length, but differ therefrom in that they are
longer— by an inch instead of shorter by that amount— than their coverts. The 1st
metacarpal remex
The
The
is distinctly
smaller than the rest of the series
ventral aspect of the wing
is
:
see p. 157.
bare.
coverts of the dorsal aspect are arranged in obliquely transverse series, the
feathers of each series being piled as it were one upon another, from six to nine
—
PHYLOGENT OF THE PAL^OGNATH^ AND NEOGNATH^.
such coverts being included in each of these bundles.
form a
series of partitions, dividing the post-axial
The
157
bases of these coverts
border of the wing into a number of
wide and deep compartments.
Two
Carjml covert and remex.
(PI,
different sets of feathers
seem
of which
me
is
XLV.
12.)
fig.
claim identification with these
to
The
invariably present in the Neognathce.
to represent the carpal
of the metacarpal series and has a large major covert.
to the
2nd remex,
The second
This
In
its
is
to
smaller than the rest
position and
its
relations
remex of many Neognathoi.
closely resembles the carpal
it
remex appears
1st metacarpal
remex of the Neognathce.
one or other
;
set of feathers, referred to as possibly representing the
Neognathine carpal
covert and remex, lie immediately pre-axiad of the base of the major covert of the
remex
1st metacarpal
or, in
;
others words, pre-axiad of the base of that featlier which
we have just tried to show may represent the carpal covert {c.c,
The identification of the covert must stand or fall by the quill.
The possible carpal remex, c'.r'., as we have just remarked,
pre-axiad of the supposed carpal covert [c.c, PI.
purposes of identification,
with the
serial
t.
medue
row of
to a transverse
we may mention
XLV.
that this
on the other.
its
relation to the 1st metacarpal
representing the true carpal remex,
between the carpal
XLV.
PI.
It
remex and
remex
c.r. fig.
the
is
12,
it
**,
seen to be
is
the terminal feather
its
covert represents
is to
say with regard
also to be regarded as possibly
closely agrees with the relation
metacarpal
remex of
Neognatlm
{c.r.,
12).
fig.
does not seem possible to settle this question definitely.
now under
two feathers
is
and
In position, that
—which
— very
1st
it
12).
immediately
lies
new remex,
In other words,
fig.
Furthermore, for the
12).
of the forearm on the one hand, and
six
the 1st median and minor covert of the manus.
to
fig.
XLV.
PI.
It
may be
that these
discussion may, in the Neognatha', represent sometimes one,
sometimes the other of these two
This seems the more probable when we
sets.
remember the varying development and
which obtain
relations to the primary remiges
in wings of Neognathce.
If the
first
correct, then
and most favoured interpretation of the identity of the carpal remex
it
would seem that
1st cubital remex, as
it
represents, in Neognathte, the 1st metacarpal, not the
was contended by Mitchell
correct, then the carpal
remex
is
of the
[62].
Neognathw
is
If the second interpretation
is
represented in Bhea by a median
and minor covert of the metacarpus.
T, majores.
On
the
— On the cubitus these are
manus they
many Neognathw
are slightly shorter.
the cubitals are at
slightly longer than their remiges,
It is
first
shorter
performing for a brief time the function of the
T. medice.
VOL.
xv^.
—The median
—PABT
V.
No.
7.
about 1
significant to note that in the
than their coverts
quills, as in the case of
;
young of
tlie
young
in.
coverts
Gallince.
coverts of the cubitus ofler no features for discussion other
December, 1900.
2a
ME. W.
158
PTCEAFT ON THE MORPHOLOGY AND
P.
than those included in the general remarks on the arrangement of the coverts.
median coverts
are only two
There
may
the hand, and, as already hinted, one of these
in
represent the carpal remex.
T. minores.
wanting
—There
in the
rows of minor coverts on the cubitus.
are 6 to 7
They
T. marginales.
—A broad
bare space separates these from the minor coverts.
are arranged along the pre-axial border of the wing, in groups of 3, just as in
They
Neognathce.
Ala
spuria.
are
manus.
They
some
are wanting in the hand.
—There
ai-e
5 thumb-quills, as in the Neognathce.
Parapteron and Hypopteron.
— These humeral remiges are wanting
in Bliea.
The feathers along the pre-axial border of the humerus are of great length.
The whole under surface of the wing is perfectly bare, a probably degenerate
character.
— There are no
Uropggium. — Absent
adult
Pt. caudce.
rectrices.
in
The Bhamphotheca,
(p. 159.)
like that of
Dromams,
sheath of the upper and lower jaws.
There
The Podofheca
of
cere,
Rhea americana
is
is
made up
of three pieces, both in the
no trace of the denticulations found in
The
or embryo.
Dromceus, either in the adult, nestling,
which are pierced through the
is
aperture of the nostrils,
lateral.
(fig.
1 d), according to Dr.
Gadow
[25], resembles
that of R. danvini, and differs from that of E. macrorhyncha in that the whole of the
acrotarsium
is
clothed
in large scutes, whilst in R.
confined to the distal extremity.
and Dromceus.
tibio-tarsus
It
is
is
R. darwinii
is
The
macrorhyncha these scutes are
leg of this last thus resembles that of Casuarius
furthermore peculiar in that the feathering of the
continued downwards on to the tarso-metatarsus for some distance.
interesting and
significant
to note,
region in the embryo of R. americana.
however, that a few feathers occur in this
Indeed, in this latter their extension
may be
downwards to within a short distance of the toes, as will be seen in fig. 2, p. 155.
The planta is also invested by large scutes.
In the f-grown nestling and adult
traced
these are arranged in the form of a series of paired plates.
embryo they form a
series of single transverse scutes.
large scutes will be found a
number of
apparently gradually increase in
size,
smaller,
wedged
and thus give
In the nestling and ripe
Along the inner
in
between the
rise to the
side of these
larger.
These
secondary arrangement
of paired plates.
The claws
of the toes are laterally compressed, and
ridge or keel, not
met with
outer and inner toes.
in other Paloeognathoe.
form a strong median dorsal
This
is
most marked
in
the
PHYLOGENY OF THE PAL^OGNATH.^ AND NEOGNATH^.
b.
The
ripe
Pterylosis
of
the
159
Embryo and Nestlmg.
embryo and the nestling Rhea americana appear
only in that the anterior portion of the apterioa sjnna/e
is
to differ
absent,
from the adult
and that there
is
a
distinct oil-gland.
I find
the egg.
no trace of an " egg-tooth
This
is
also
wanting
"
Concerning Uroincens, however, see
find
it
in
Apteryx
at
any stage of
in the ripe embryo,
which
the embryo Dromceus and
in
p. 154.
its
The
I
myself removed from
the nestling Casuarius.
Prof. Jeffery
late
Parker failed to
growth.
STRUTHIONID.iE.
Struthio camelus.
1
have not been fortunate enough
consequently
I
am
obliged to
fall
to secure
either nestling or adults of Struthio
back upon the published accounts of others.
;
Nearly
Fig. 3.
B.
A..
Lateral view of an embryo of Struthio, after Lindsay, to compare with Apterijx.
Note the absence of a
lateral cervical aptcrion.
B.
Lateral view of an embryo of Apteryx
the distinct pieryla
all
colli
ventralis
austral'ts, right side (after
and dorsalis and the pteryla
Parker), to
show the
pterylosis.
Mote
humei-alis.
that can be gathered, moreover, concerning the pterylosis of the trunk of Struthio
concerns embryos only.
The wing
of the adult Ostrich has been ably described by
2a2
MR. W.
160
Wray
[97]
;
PTCEAFT ON THE MORPHOLOGY AND
P.
the figure illustrating his paper has been reproduced here by the kind
permission of the Zoological Society.
been figured by Lindsay [48], and
Apteria
Apt. spinale.
is
The
pterylosis of the trunk of the
reproduced here
—
than
— Very narrow, longer
Hildebrand
—According
[37]
relatively
Apt. mesogastrcei,
cloaca, but judging
to
(fig.
3a).
in Casuarius,
this extends
from Miss Lindsay's [48] figures
embryo has
which
it
resembles.
from the beak to the
did not extend beyond the
it
sternal callosity, as in other flightless Palceognathce-
Apt. trunci laterale.
of Apteryx, and
—This would
to differ
from
all
appear in
and form
size
to closely resemble that
the remaining PalmognatluB.
Hildebrand also describes an apterion on the crown of the head.
Pterylce
—
Ft. alaris (PI.
digitals,
XLV.
—According
—metacarpals
11).
fig.
distributed as follows
:
Wray
to
8
;
[97] there are 16 metacarpo-
addigital
1
mid-digitals 4
;
;
pre-
digitals 3.
According to Wray's
figure,
reproduced on PL
XLV.
fig.
11, the
primaries are
placed wide apart and almost at right angles to the long axis of the hand.
number of the metacarpals
Neognatlue,
i.
e.
is
higher than in any other bird
Flamingo, there are 7 metacarpals.
—
8.
In
The
Bhea and some
In no other bird, save Struthlo,
The arrangement of the coverts appears
The ala spuria is made up of 4 remiges.
are there 4 mid-digitals and 3 pre-digitals.
to
differ materially
from that of Ehea.
before attempting the search for the carpal remex and
I await a spirit-specimen
its
covert.
It
would be interesting
to
know whether the^cror
carpi ulnaris resembled that of
Bhea in the large fleshy portion for the attachment of the remiges (fig. 5 a, p. 240)
and what is the condition of the vinculum tendinese. Assuming that the disposition of
these parts in Rhea represents a proto-carinate condition, we should expect to find the
wing of Struthio very similar in these particulars.
Both poUex and index are armed with a claw.
Mhamphotheca.
Podotheca.
scutes.
— Compound
—Acropodium
Only 3rd and 4th
a claw.
Uropygiutn wanting.
as in
Bhea
and lower portion of acrotarsium with broad transverse
toes are present
;
of these in the adult only the 3rd bears
::
PHYLOGENT OF THE PAL^OGNATH.E AND NEOGNATH.E.
IGl
Apterygid^.
Aptertx australis mantelli.
—
Apteria
A. spinale.
A.
—This wanting.
—This a well-marked
is
inesogastrcei.
space, extending forwards from the cloacal
is
aperture, along the middle line, as far as the base of the neck, to a point corresponding
to
the interclavicular region of the Neognathce.
the
abdomen
middle of the sternum
in the
;
A. trunci laterali
is
it is
Its greatest
width
in.)
(1
on
is
nearly ^ in.
It runs
of very considerable extent.
from a point immediately
over the head of the femur downwards and forwards as a conspicuously broad bare
space of about 1
in. in
gives off a branch
width
;
terminating at the base of the neck.
At
the wing
it
downwards and backwards between the thigh and the pteryla
ventralis as usual, and terminates about 2^ in. from the cloacal aperture.
—
Pteryla'
Pt. capitis.
—This very dense, and bears numerous
—Forms a broad saddle-shaped
is
tract,
and anteriorly
Pt. ventralis
and
is
is
of the
Pt.
passinij into the
Anteriorly
well defined.
j^t- colli.
blends with an incipient
it
tlie
'pt.
colli ventralis,
other by the apt. mesogastrmi.
pt. colli is almost divided into a dorsal
ap)t.
;
Yet
XLV.
fig.
addigital 1
The innermost
position.
and ventral
tract
by the forward extension
trunci laterale.
alaris (PI.
metacarpals 1
I
metacarpal remex
cubital
is
13).^Rennges
mid-digitals
;
13.
(\
very weak, and
mid-digital)
is
is
first quills
Metacarpo-digitals 4
9.
—
2.
is
counted a remex solely on account of
cramped in
me
position,
The calamus
portion to the length of the whole feather, and
these quills
Cubitals
think that most would agree with
into the position of a major covert.
the
the
continued backwards as a sharply defined tract bounded on the one side by the
apterion trunci laterale and
The
blending posteriorly with
tract
Pt. spinalis.
femoral
rictal bristles of great length.
in
this
its
The 3rd
and forced dorsalwards almost
of these quills
is
decision.
is
very long in pro-
fusiform in shape.
exactly reproduced, as will be seen in the figure (PI.
The form
XLV.
fig.
of
15), in
of the nestling Cassowary (see p. 165).
The proximal end of the forearm, over the region of the elbow-joint, is bare, as in
many Neognathce.
The arrangement of the coverts is that of obliquely-transverse rows, as in iV^ognathw. Only tectrices majores, however, can be certainly made out. There appears
to
be but one major covert on the manus.
The
index-finger bears a large claw.
Pt. femoralis.
point.
— Uniformly
There
is
no pollex.
feathered, passing forwards
into
a sharp, cone-shaped
ME. W.
162
The Uropygiuvi
is
PTCRAFT ON THE MOEPHOLOGT AND
P.
a large, compressed, naked, conical gland lying at the bottom of a
conspicuous depression
— formed
by the great development of the gluteal muscles,
and immediately above the cloacal aperture.
Mr. Beddard's recent paper
in
Bhmnphotheca.
gnathce
;
but
this is
extreme
masked by the
but this
tip,
Podotheca
where they
will be
found
up of the same elements
as in the other Palcpo-
greatly elongated form of the beak.
There
is
a
not pierced, as in Rhea, Dromceus, and Sfruthio, by
is
(fig.
2 e).
found as a pair of minute
will be
—The
scales, or of large scutes as in
slit-like apertures.
may be either made up of large rounded
The latter form occurs, so far as I
A. australis mantelli. The planta in this species
acrotarsium
Rhea americana.
have been able to make out, only in
is
uropygium
These, in Casuarius, open near the top of the beak, and in Apteryx at
the nostrils.
its
figure of the
[8].
—Apparently made
distinct cere at its base,
A
In an embryo of A. australis mantelli the fusion
covered with large coarse papillae.
of small plates into scutes in the acrotarsium
is
well
shown
(fig. 7).
Crtpturi.
The pterylography of
this
siderable detail by myself [8
marked than
in
1].
group of the Palceognathm has been described in conIt will suffice
here to say that the apteria are more
In the form of the apt. spinale, Calodromas
the flightless forms.
elegans, one of the Crypturi, closely resembles Rhea.
the group, filo-plumes occur
— though
however, are restricted to the pteryla
The
forms
;
Unlike the
very sparingly; and down-feathers.
dift'er
tomium
These
from those of the
but in the remarkable development of the radii and
denticulated
members of
last,
alaris.
neossoptiles, in the large size of the aftershaft,
in the
shape of the rami, the Crypturi closely resemble the Struthiones
The
flightless
flightless
elongated flattened
(PI.
XLV.
fig. 2).
of the nestling Calodromas recalls that of Dromceus.
The Structure of the Feathers.
The Neossoptiles.
The neossoptiles of the Palceognathxe consist of prepennae only, but these present
many features of great interest.
The prepennae of Casuarius have a moderately long rhachis, bearing some 3-4 pairs
The radii are unsegmented, of relatively medium length, and bear minute
of rami.
The aftershaft is apparently represented
fila, sparsely distributed (PL XLV, fig. 5).
by about 6 rami standing at the base of the main
rhachis, as in the
'J'he
Fowl
or
shaft,
and not seated along a separate
Tinamou.
prepennae of Promcens (PL
XLV.
figs. 4,
4 a) appear to be distinguishable from
those of Casuarius chiefly, and probably only, on account of their rather greater leugth
;
PHYLOGENT OP THE PAL^OGNATH^ AND NEOGNATH^.
The
the rhachis bears about 6 or 7 pairs of rami.
As in Casuarius, fila are minute and rare.
The nestling-down of Aptenjx (PI. XLV.
of the remaining Palceognatha;.
well-developed shaft, but
numerous rami.
From
it differs
163
radii are slightly stouter, relatively.
from that
6) is easily distinguishable
fig.
Like that of Dromwus and Casuarius, it has a
from these in its greater length and in the very
EJiea and Struthio
entirely, in
differs
it
that, in these, the
neossoptiles are umbelliform.
striking feature perhaps of the nestling-plumage of Apfertjx
The most
length.
texture
In
we appear
to
get three gradations.
australis very closely resemble one another
and
differ
is
its
great
A. australis mantel li and
from the remaining species
in
A. oiveni and A. haasti
that the feathers are coarser and slightly harsh to the touch.
Some individuals of A. australis,
stand at the other extreme and are peculiarly soft.
however, seem to stand between the two.
A. haasti
is
the most markedly distinct of
all.
The
feathers clothing the head, neck,
texture, the down-feathers
and thighs are of a peculiarly downy character, recalling, in
breast they are of a peculiar
on the breast of tlie adult Duck. On the back and
resembling very strongly the intermediate plumage or pseudo-
"woolly"
texture,
definitive feathers, (mesophyles of Palmer),
There
is
no trace of an aftershatt
which follow the prepenn«
in the Owls.
to these prepennae.
20 pairs), and moderately long. The
of being formed of a number of
radii are relatively short, and have the appearance
of
compressed joints, the dorsal and ventral angles of the anterior ends
The rami
are closely set, very
numerous (18
to
short, laterally
which are produced forwards to form a pair of minute lila.
the rami tlie prepenna of
In the length of the shaft and the great number of
it is further peculiar in that
stands alone amongst the Pala^ognathw; moreover
produced beyond the most
A
distal radii into
ylpteri/x
it is
not
long filaments.
peculiarly interesting feature of the prepemue,
and of the
first definitive
feathers,
in all other
they appear to do
that the latter do not thrust out the former, as
investigation.
Exactly how this is avoided is a point for further
birds.
known
is
I
hope
to
be
That this process of thrusting-out is avoided
able to explain this mystery shortly \
Here
haastii in the Kothschild Collection.
can be still seen in a downy nestling of A.
about halfway through the skin, and show no trace
other Palm- and NeocjnatJm.
whatever of a prepenna surraountmg the tips, as do the
the sheath common to
Furthermore, they are peculiar in that they are not invested by
the
all
first
definitive feathers are
developing feathers.
This appears to break away as soon as
it
reaches the surface
of the skin.
plumage may represent preplumoe instead of prepennie
in the
hy the teleoptiles. The fact that plumula; do not occur
this would account for their non-ejectment
but
Ukea,
embryo
"
described " pinsel-artig down-feathers in the
adult is of no importance. Zander [loi] has
'
It
may
possibly turn out that this nestling-down
this is probably
an error.
:
ME. W.
164
The
PTCEAFT ON THE MOEPHOLOGT AND
much from
structure of the adult contour-feather does not differ
The
prepenna.
be in this
the shaft
sliaft in
:
chief distinction
—that
;
—apart from
in the adult the
size
—whereby the two
rami are arranged in
pairs,
that of the
differ
appears to
and not alternately up
that the radii are confined solely to the rami, and do not grow along the
the inter-ramal spaces
nodes and internodes
the
P.
;
;
and, thirdly, the rami are not so distinctly divisible into
but there
is
not
much
difference in the form
and length of
fila.
Casuarius,
main
shaft,
Drommis, and Apteryx
and no
all
agree in that the prepenna has an elongated
distinct aftershaft.
Bhea and Struthio agree one with another, and
that the prepenna
is
umbelliform (PI.
XLV.
differ
figs. 3,
from the above-mentioned, in
The main
3 a).
shaft appears to
be represented by 3 rami conspicuously stronger than the rest with which they are
The prepennse are borne upon the tips of the definitive feathers for a very
associated.
considerable time, giving a quite peculiar appearance to the first plumage.
In Rhea the bases of
all
the rami, 9 in number, are seated around the upper
—
feather— the rhachis as just remarked, is represented
by 3 elongated rami, bearing radii from the base upwards for a short distance, and
produced beyond these into a long slender filament. The aftershaft is represented by
umbilicus.
The main
axis of the
some half-dozen rami, furnished
with
numerous
delicate
radii.
These
last
are
moderately long, strap-shaped filaments, with scarcely the faintest trace of fila.
In Struthio the typical neossoptile (PI. XLV. figs, 7, 7 a) has the three rami representing the main shaft produced far outward beyond the tip of the feather
projecting region of each
lamina, liollowed in
its
taking the
;
the
form of a hardened, glistening, ribbon-shaped
ventral surface.
In this particular
it
closely agrees with the
and run along the rami as a series of relatively broad,
flattened, and pigmented laminae, set obliquely on to the ramus, and overlapping one
Moreover, each
another much as in the normal adult Neognathce contour-feather.
Crypturi.
The
radius bears
radii are short,
numerous but
and booklets.
The forma-
many Neognathw
are worthy
small, probably degenerate, fila
" eye-spots " seen in these radii of
tion of the remarkable
of notice and further examination.
The
aftershaft rivals the
not, however, differ
main
shaft in size,
from those of the main
numbering some 12-15 rami, which do
shaft, save only in
that the rami are not
produced into elongated horny bands, whilst the radii are larger than those of the three
rami representing the main shaft.
165
PHTLOGENY OF THE PALJiOGNATH^ AND NEOGNATHiE.
The Structure and Homologies of the Adult and Nestling
Eemiges of Casuarius.
The
Nc.dliiiq.
—The
youngest nestling in which these can be studied
is
that of a
stuffed C. casuarius salvadorii (Rothschild Mus.).
In the dried wing of this there
remiges.
are 6
as in Neognathoi.
wing-membrane
seated in a distinct posterior
These are widely spaced, and
Each
consists
of a
moderately long calamus and a long and tapering rhachis bearing from 4-5 pairs of
symmetrically disposed rami set very far apart. There is but the feeblest trace of the
fusion between the prepenna and the displacing definitive feather
In
The
now under
discussion.
casuarius sclateri the growth of the wing-spines has proceeded a stage farther.
C.
definitive feather
is
seen to bear rami as well as prepennse.
prepenna.
In Casuarius loriw the quill-spines have grown very long and have lost the
rami.
few
a
bears
yet
The 5th or innermost spine, representing the 3rd cubital remex,
In this stage, and in the
last described (C.
c.
sclateri),
we
get an inkling into the
developmental history of these remarkable wing-spines.
feather,
This is not the place for a recapitulation of the developmental history of a
yet
it
would be well
standing of
what
to recall one or
is to
facts
concerned with this for the better under-
follow.
axis of a feather, then,
The
two
is
two parts— the calamus or
divisible into
quill,
and the
of the dorsal region
rhachis or shaft. The latter appears to be a continuation forwards
This being so, it is obvious that this drawn-out portion must
of the tubular calamus.
These in the normal feather turn inwards towards the
free edges.
have two lateral
middle line and
meet, leaving a very fine seam to indicate the line of junction.
will be remembered, is a thin, dense, horny, and semitransparent
finally
But the calamus,
it
structure and perfectly hollow, whilst the rhachis in section
is solid,
being composed of
medullary portion or
an outer layer continuous with that of the calamus, and an inner,
to be
pith— a white, soft, cellular structure resembling elder-pith. This pith seems
horny membrane, thus insuring lightness,
for the sake of supporting the outer
added
toughness, and strength.
The
The whole forms
the shaft.
dorsal elongated region of the shaft,
now
swollen out by pith, forms a sort of
The ventral edge of this mouth
plug to the otherwise open mouth of the calamus.
umbilicus.
always visible. It forms the boundary of the upper
The
is
of the Cassowaries, represents
aftershaft, so conspicuous a feature in the feathers
whicli obtains
of the ventral region of the calamus exactly similar to that
an elongation
on the dorsal. Thus we have a dorsal and ventral rhachis.
have only the dorsal rhachis, and
In the remiges of C. lorice and C. c. sclateri we
being no pith-cells.
great interest as only its outer layer is present, there
this is of
ribbon-shaped lamina whose free
This layer takes the form of a delicate scroll-like,
material, or pith, they fail to
edges turn inwards, but from the absence of the packing
meet
in the
VOL. XV.
middle line
— PART
V.
(PI.
XLV.
fig.
14
a).
No. S.— December, 1900.
2 b
ME, W,
16C)
The
fate of the
P.
PTCEAFT ON THE MORPHOLOGY AND
prepenna which remains attached
rami appears
to
from the calamus by which
it
definitive rhachis
to
break
off
with
its
its
so familiar in,
to,
differ,
rhachis and
Carinatm,
The
fracture takes place at
This appears to go on growing and
it
becomes the
solid, elastic spine
the Cassowaries.
In the wing of the adult A-pteryx
These
The
was supported.
walls, so that in course of a short time
and peculiar
off as in
be carried for a short while longer, and then
the upper umbilicus, and leaves only the calamus.
thickening
rubbed
It is shed or
be conjectured with probable certainty.
the young definitive feather can
to
we have an almost
similar series of wing-spines.
apparently, from those of Casuarius only in that they have retained the
rami, and in that the calamus
its
is
In one point, however,
yet hollow.
they differ markedly, inasmuch as they have what
is
possibly a vestigial aftershaft.
This can be readily seen as an elongation of the central
lip of
the calamus, the sides
of which bear numerous rami.
The
the
In
Palceognathe there
is
Teleoptiles.
but one form of teleoptile, corresponding to the
contour-feathers of the INeopiatlue, semiplumulse, pluraulse, filoplumulae, and powder-
down
as are
Rictal bristles, such as occur in Ajjteryx, and eye-lashes, such
are all absent.
found in
all
other Palceognatlm save A^teryx, are modifications of contour-
feathers.
Meijere claims to have discovered filoplumae in Struthio, Rhea, and Casuarius, and
gives figures of each.
feathers
in
paimanus
C.
the
:
I
cannot, however, confirm this.
wing of Casuarius similar
that
have found degenerate
he figures from the neck of
these, however, I regard as degenerate contour-feathers.
anything in Rhea corresponding to his figure
of examining.
He
failed to find
In Casuarius the teleoptiles
and
to
I
them
are,
radii,
Struthio I have not had an opportunity
in other Palceognathoe.
on the lower region of the back, of great length,
relatively larger than in Bromceus.
from wherever taken, bear
;
I failed to find
Only the rami of the lower part of the
and these appear
to
be entirely destitute of
feather,
fila.
The
rami throughout the greater part of the length of the feathers are long and hair-like,
reminding one of egret-plumes.
The hyporhachis
is
nearly or quite as long as the main shaft, and does not differ
therefrom structurally, as in Neognathoe.
The Casuariidm
borne by the wing.
The plumage
its
are remarkable for the
spines, 5 in
number, which are
These are degenerate remiges, and are dealt with above.
of the Dromceidce differs conspicuously from that of the Casuariidm by
greater softness of texture.
The rami
enormous
In the latter
it is
harsh and coarse.
are closely set, as in typical contour-feathers of Neognathce.
In a feather
taken from the middle of the pteryla spinalis the radii are of great length, and give
PHTLOGENT OF THE PAL^OGNATIL^ AND NEOGNATH.E,
the surface of the feather a very
and
in lengtli,
whole length
—
is
distal
laminate mass.
here and there under the microscope.
only minute pointed
On
appearance
fila,
for the
;
towards the
end of the feather
peculiar, in that the rami bear
common
pletely into one
soft, silky
The
finally disappear.
no
radii,
167
tip they decrease
— about one-sixth of
but fuse more or
less
its
com-
Separate rami may, however, be distinguished
Tlie radii are very long, filamentous, and bear
most part unpaired.
the neck the feathers are of a looser structure, the radii being shorter, and
disappearing
The
much
sooner than in the back-feathers.
contour-feathers of the Apterygidce appear at
those of
many
fact that
it is
first
sight to differ but little from
Neognathce, the vane of the feathers, seemingly, being continuous.
discontinuous
is
The
only proved after microscopic examination.
The rami are paired, set close together, and moderately long. The radii are long,
filiform, more or less distinctly jointed, and armed with numerous minute fila, set in
pairs, following close
one upon another.
These run
in this fashion
through almost the
entire length of each radius, only the extreme proximal end lacking such appendages.
According to Parker [71] tliey are longer in the embryo than in the adult. The entire
absence of booklets at once distinguishes this from a laminate feather.
There
is
no
(See also
aftershaft.
p.
163.)
In the Stnithionidw the adult contour-feather more nearly resembles that of the
Neognathxe than
The rami
is
the case with any other of the Palceognathw.
The
are set closely together, precisely as in the typical Carinate feather.
radii take the
form of narrow blade-shaped bands, directed upwards and outwards, so
that the rami from which they spring form the bottoms of V-shaped troughs.
distal
end of each radius bears
sufficiently elongated to
Moreover, the radii
radii
shaped.
Thus, the vane
:
they are probably, however, vestiges of these.
from those of more perfect feathers
differ
is
The
disposed as in normal Neognatlue, but none are
form booklets
differ
do not
proximal
fila
one from another
discontinuous
:
in that the distal
and
both series are alike, long and blade-
— a probably degenerate condition.
In the Mheidce the contour-feathers of the trunk are characterized by the markedly
discontinuous character of the vane
between which
;
the rami appearing as a
number of hard
The remiges are of very considerable size, though far inferior to
The rami bear numerous short and fairly conspicuous radii.
The radii of the track-feathers, under the microscope, appear
rather less than have those of the remiges.
proximal end, each radius
its
4 or 5 long
fila,
lines,
radii are only faintly traceable.
is
those of the Ostrich.
to
have degenerated
At
lamina bearing some
They approach those of the Ostrich.
laterally compressed, the resultant
probably vestiges of booklets, and of the processes which occur on the
ventral edge of the lamina in the proximal radii of Neognathine feathers.
The
fila
radii of the
remiges are
suggest vestiges of booklets
filiform,
;
but distinctly flattened from side to
they are frequently
side.
bifid.
2b2
The
ME. W.
168
P.
PTCEAFT ON THE MORPHOLOGY AND
The
In the DmornifhidcE the rami and radii are short.
the feather to
its
base, are filiform
and divided
latter,
into a series of
from the middle of
numerous short
the anterior ends of which are produced into two or three pointed
radii
most nearly approach those of Apteryx, but they may be
therefrom by the relatively great length of the
The
distal
The
ends of the rami bear no
Dinorms were
feathers of
appended below
"The
fila.
radii.
described by Dallas [15].
first
His description
web
is
somewhat
different
from that which occurs
Towards the base of the shaft the barbs spring
and the Cassowary.
or five together from nearly the
number
Thus these
fila.
once distinguished
is
:
structure of the
tufted aspect.
at
joints,
same
spot,
and thus
As we advance towards the apex
this
in the
Emu
groups of four
in
web assumes
this part of the
arrangement speedily ceases
;
a
the
of barbs springing from the shaft gradually diminislies until each side bears
The barbs
only a single series of these appendages.
consist of slender, flattened fibres,
bearing long, silky, and very delicate barbules, without any trace of barbicels, and
presenting a distinctly beaded appearance
into a series of cells,
some of which towards the apex exhibit small tooth-like projections
The barbs nearest the base
representing rudiments of barbicels
the main
web and
barbules throughout
its
of
is
feather,
tlie
some distance
accessory plume, are destitute of barbules for
from their base; but this distance gradually decreases until the barb
As
Under
a simple lens.
the barbule appears merely divided by faint transverse partitions
the microscope
botli in
when examined by
furnished with
whole length."
Dallas remarks, these fragments
still
leave us in ignorance of
many
such as whether the basal rami (barbs) possessed the hair-like tips of the
points,
Emu
and
Cassowary, and whether the apical portion of the feather supported simple rami as
in these last.
SUMMAEY OF PtERYLOLOGICAL CHARACTERS.
Perhaps the most interesting pterylological result of the present enquiry has been
the light thrown
of Casuarms.
upon the
history of the remarkable wing-spines or vestigial remiges
The comparison
of these with the remiges of the nestling on the one
hand, and with the wing of Apferyx on the other, leaves
little
or
no dmibt but that
the explanation, or rather interpretation, of the nature of these degenerate feathers
is
correct.
The form and
disposition of the apteria have been described at greater length than
ever before, and some
at least in Struthio
new
facts added.
The
existence of apteria has
and Apteryx; though these
the text-books, which
still
facts
kmg
had not yet found
been known
thiir
way
into
repeat the old error that the feathers of the FcdoeoguathcB
weie evenly distributed over the body.
The form and
structure of the nestling-down has been
worked out
iu considerable
!
169
PHYLOGENY OP THE PAL.EOGNATH* AND NEOGNATH^.
and
in the complexity of the
main
shaft,
An
come about
(PI.
XLV.
fig.
13).
Casuarius, by a few sessile rami.
sessile
how
rami without any trace
the loss of this
the aftershaft
:
umbelliform, and
is
is
may have
represented, as in
In Struthio, however, the number of these
In both Bhea and Struthio the main
in the latter are
In these particulars,
shaft.
In Rhea and Struthio the prepenna
thereby differs from that of the other Palceognatlue
which
few
of the adult Aptenjx show
The remiges
of a rhachis.
there a distinct after-
aftershaft appears to be represented
here it consists only of a
only in the two former, and
is
In Casuarius, Dromceus, and
unique.
is
it
Apteryx, these have a well-developed vhachis.
great.
main
this is of great size, equal, indeed, to the
and
shaft,
In the Crypturi only
Prepennae only are represented.
detail.
sliaft is
very
is
represented by 3 thickened rami,
produced forwards beyond the
rest of the feather to
form broad,
hollow, ribbon-shaped lamime, recalling in form the nestling-dowu of the Crypturi
(PI.
XLV.
fig. 2).
The following point involves a mystery whicli I am anxiously endeavouring to
The prepennae are regarded by some as nothing move than portions of the
extremities of the developing rami of the teleoptiles below.
it
If this
is so,
solve.
distal
how comes
that the prepennae of Casuarius and Dromceus have a scarcely recognizable aftershaft,
whilst in the teleoptile
the main shaft
A
is
of such great length as to be hardly distinguishable from
Tinamous these
In the
1
in tlie adult feather
main
it is
The
relations are exactly reversed.
very small or wanting, and
in
the nestling
it is
aftershaft
as long as the
shaft
further most serious objection to the probability of the truth of this view
fact that in
These
Apteryx the nestling-down feathers are not driven out by the
arise at
nestlings
is
the side of the prepennae,
not large enough to show.
definitive feathers
It
the
is
teleoptiles.
my
series of
probable that they are shed as soon as the
have completed their growth.
feathers does not seem
ultimate fate of which
the
is
The
peculiar
downy nature
of these
to have been recorded before.
The discovery of a uropygium in Dromceus and Mliea is a point of some interest.
The podotheca appears to be of some slight value for systematic purposes. Dr. Gadow
I have
[25] long since pointed out the difierences between the three species of Bhea.
of
different
species
between
the
not succeeded in finding any appreciable difference
Cassowary, or in distinguishing that of Casuarius from Dromwus.
seems
(fig.
2
to differ
B,
p.
from the other species in
155), just as
this respect
and
Apteryx australis mcmtelli seems
to
Casuarius
form a type of
to
differ
lorice
its
own
from the other
Apteryges in having, as a rule, the acrotarsium clothed with transverse scutes in place
of small rounded plates.
Dromceus
the
represents
undoubtedly comes next
;
less
specialized
of
all
the
tliough in the brilliant coloration of
remarkable casque, spine-b'ke remiges, and elongated claw on
a distinct advance upon Dromceus.
Casuarius
Palceocjnatho'.
its
its
head and neck,
inner toe
it
has
its
made
ME. W.
170
PTCEAFT ON THE MOEPHOLOGY AND
P.
Apteryx should be placed next.
It
has
In the large
prepenna has a well-developed rhachis.
possession of an oil-gland
the aftershaft to the feathers
lost
resembles the Carinatce
it
size of the
so also
;
the
;
apteria and in the
does in the softness of
it
the neossoptiles.
Ehea and Struthio are undoubtedly the most highly
of this, from the point of view
of the chief pieces of evidence
characters,
upon that of Casuarius or
The
fact
two than in the remainder of the group, only means
larger in these
is
that the need for this organ remained longer, and
wing
of pterylological
the fact that the prepennae has neither aftershaft nor rhachis.
is
that the wing
One
specialized of the group.
is
not to be regarded as an advance
may
Apterycc, as the case
be.
In them the decline of the
set in earlier.
The
groups
structure of the teleoptiles in these last two
:
is
more complex than
These have lingered longer, just
booklets are feebly represented.
in the other
as the
power
of flight.
We
may
as follows
briefly
sum up
the salient features of the pterylosis of the PalceognatJm
:
All the flightless Palceoguathce agree, and differ from the Neognathce,
the under surface of the wing naked
^,
and
in having
have discontinuous
in that all the feathers
vanes.
A. Adults.
Lacking a pollex.
DronuEUS, Casuarius, aud Apteryx have ao ala spuria.
The
teleoptiles of
Dromaus and Casuarius have
a large aftershaft.
There
DromcBHS has numerous though greatly degenerate remiges
Casuarius has few remiges
to
form
elongated
-6
in
all.
— 17
is
no oil-gland.
in all.
These are degenerate and peculiarly modified
porcupine-like quills, representing only the calamus of
ordinary feathers.
The
teleoptiles of
13 in
all
Apferyx have no aftershaft aud a large oil-gland;
—aud
these have a long calamus.
it
Apteria are larger.
has few remiges
—
Rictal bristles are of
great leugth.
Rhea and Struthio have a large ala spuria.
The teleoptiles of Rhea and Struthio have no
numerous and
aftershaft
:
there
is
no oil-gland.
Remiges are
large.
The remiges of Rhea number 28; metacarpo-digitals 12. There are no rectrices.
Struthio number 32 metacarpo-digitals 16.
There are large rectrices.
The remiges of
;
Apteryx aud the Crypturi are the only Palseognathinse lacking a sternal callosity.
'
The naked under
surface of the
to the suppression of the feathers
wing
of the flightless Struthious birds is probably a secondary feature
by the long continued application of the wing to the
side of the body.
due
;
PHYLOGENY OF THE PAL.^OGNATH.E AND NEOGNATH^.
171
B. Nestlings.
The
neossoptiles consist of prepennse only.
In Dromceus, Casuarius, and Apteryx the prepenua has a well-marked rhachis and numerous
rami.
The
aftershaft
is
feeble or wanting.
In Dromeeus the rhachis bears about 8 pairs of rami
:
the rami are produced into
points beyond the distal radii.
In Casuarius the rhachis does not bear more than 5 pairs of
In Apteryx the rami exceed 8 pairs
;
the prepennse of some parts recall the adult
In Rhea and Struthio the prepenua
is
radii.
the rami are not produced beyond the radii
umbelliform
;
down
the main
of Ducks.
represented by 3 rami,
sliaft is
the aftershaft by numerous rami.
In Tihea the rami of the main shaft are not greatly elongated or
In StrutMo the
ends of the rami of the main shaft are
distal
and grooved,
peculiarly flattened
In Crypturi the prepenna
is
very complex
;
flattened.
much
elongated and
as in Tinamidce.
the aftershaft equals the
main
shaft in length.
OSTEOLOGY.
The Skull of the Adult.
The
skull of the Palmognatluv
and Tinamous
so-called Ratitce
—
the pterygo-palatine connection
The
— which, as
we have elsewhere shown, includes the
from all other known birds in that, in the adult,
differs
is
by symphysis or anchylosis, and not by an arthrosis.
sknll of the Tinamous, as will be
shown
presently, closely resembles that of
—The
occipital condyle
Bliea.
The Occipital Region (PL XLII.
plane of the occipital foramen
fig. 7).
inclined backwards, in
is
Dinornis, yUpyornis, and Aptergx, and in the Tinamous.
occipital
foramen
is
nearly vertical
In Dinornis and ^pyornis
this
;
in
all
is
sessile,
and the
the Paloeognathce save
In Dinornis the plane of the
Apteryx the occipital condyle
is
pedunculated.
condyle projects beyond the dorsal margin of the
foramen.
In Dromeeus and Apteryx the supra-occipital, immediately above the foramen
magnum,
presents the concavo-convex
Tuhinares and Sphenisci.
There
is
" cerebellar
dome
"
so
characteristic
a well-marked snpra-foraminal ridge (PL
of the
XLII.
fig. 8) running laterad on either side to become lost on the paroccipital process in all
In Apteryx the ridge ends abruptly in the form of
save Apteryx and the Tinamous.
two pendent tubercles on either
Tinamous
side of the
the ridge terminates in the
occipital crest
is
and downwards
most strongly marked
foramen
same
in
at
about halfway down.
region, but
Rhea.
is
without tubercles.
The lambdoidal
or less sinuous outline,
and
is
lambdoidal ridge delaminate, as
not so easily followed.
it
The
ridge nins outwards
as a sharply-defined ridge, terminating in a pair of strongly
paroccipital processes, in all save Dinoriuscind Struthio.
In the
compressed
In these the ridge has a more
Only in Dinornis does the
were, in the region above the occipital crest, to
form anterior and posterior lambdoidal ridges.
ME. W.
172
The paroccifiHal
PTCRAFT ON THE MOEPHOLOGY AND
P.
processes pass gently forwards at their base into the zygomatic
process, in all save in the
Tinamous and Apteryx, Bhea, and Dinornis.
In Apteryx and
more
the Tinamous they are separated from the zygomatic process by a notch
or less
In Linornis they pass forward into the squamosal prominence, and there lose
deep.
In Ehea only, externally, they are continued upwards as a strong ridge to
tliemselves.
join the posterior boundary of the temporal fossa, and, internally, sweep round towards
the middle line in the form of a thin, curved, laminated plate of bone to form the inner
boundary of the tympanic
The Roof of
recess.
Cranium
the
(PI.
laterally, in front of the
produced
figs.
temporal
and the
processes, in all save Apteryx
all
1-4).
—The
the cranium
roof of
fossa, into a pair of
pendent post-orbital
The
reduced to a mere prickle.
save Dinornis, JEpyornis, Apteryx, and Crypturi,
terized by a pair of very long outstanding
is
anterior
charac-
and backwardly-directed spines, being the
In Struthio these are continued
elongated supra-orbital process of the lachrymal.
backwards, by means of fused supra-orbital
is
In the former, post-orbital processes
Cri/pturi.
in the latter they are
are entirely absent, and
region of this roof, in
XLII.
the frontals
ossicles, to join
space between the ossicles and the combined frontal and nasals
;
thus enclosing a
(fig. 8).
In Dinornis the lachrymal passes backwards insensibly into the frontal to form a
broad overhanging ledge to the
either side of the skull, but
is
In the Crypturi
orbit.
it
projects conspicuously on
In Apteryx
without the backwardly-projectiug spines.
the lachrymal has become reduced to a mere vestige fused with the alinasal.
In
all
save Apteryx and the
overhanging pent-house
inwards
in
Tinamous the
frontal
;
and in the Tinamous this region
almost to the middle
line, so that the interorbital
Young
and Dromceus resemble the
skulls of
in
is
Bhea
a conspicuous cerebral
Dromceus
dome
ater, the skull slopes rapidly
in yEjii/oi'nis the gradient is
The
produced laterally
to
form an
the form of a smooth convex infolding of the bone, leaving the orbit without
any sharply defined boundary
There
is
In Apteryx, the frontal passes downwards and
to the orbit.
iuter-orbital region in
much
less
Dromceus
in
is
deeply hollowed out,
region of the skull
very narrow.
is
Tinamous in this respect.
^Epiyornis and Dromceus ater.
downwards and forwards
In
in front of this
profile,
dome
marked.
is
very wide, in
Bhea very narrow
:
thus, taken
in connecticm with the development of the supra-orbital processes of the lachrymals,
this
forms a useful character for systematic purposes.
In the Tinamidce the frontals bear shallow supra-orbital grooves.
The Base of the Skull (PI. XLII. figs. 5-8). Casuarius, Dromceus, Struthio, and the
Tinamidce all agree, more or less closely, in the form of the basitemporal platform. In
all,
viewed below,
it
takes the form of a flattened, slightly tumid area, roughly penta-
gonal in shape, the base being caudad.
Its postero-lateral angles are not
produced into
mammillary processes, nor are there, save the Eustachian apertures, any sharply defined
boundaries forwards.
Struthio and Dromceus
somewhat
closely resemble one another
173
phylogelvt of the pal^.ognath^ and neognath^.
in the
form of
In
this platform.
Dmnmis
a steep face to the pre-condylar fossa
this face slopes gently
very sharply defined, presenting
in the other species of
upwards and backwards.
Dromwm
and mStrutho
Thus, in all save D. ater, the pre-
shallow and moderately wide.
and RJwa.
The pre- temporal platform is very sharply defined in Avteryx, BinornithidcB,
roughly triangular in form,
In Apteryx and DinornUhince amongst the Moas, it is
moderately or very large
postero-lateral angles are produced downwards into
condylar fossa
its
;
(iter it is
mammillary
line,
is
processes.^
and present a deep
These
in
vertical
some of the Dinornithinw coalesce in the middle
In the
face when tlie skull is seen from behind.
defined along
Emeiwv the platform takes a laterally expanded pentagonal form, sharply
ridge helping to form the Eustachian groove.
its antero-lateral angles by a raised
In Mpijorms the
The anterior basicranial fontanelle is marked by a deep groove.
The mammillary tuberobasitemporal platform is mucli shortened antero-posteriorly.
ridge along the whole of each postero-lateral side
sities run in the form of a low rougli
of the pentagon.
steep
has the shape of an elongated pentagon presentmg
The
slope gradually into the rostrum.
faces on all sides, though anteriorly it may
the basipterygoid processes.
Eustachian apertures lie immediately below the bases of
In Ehea
^the platform
other in the middle
grooves are short, widely separated one from the
In these last the
Casuarins, arched over by bone.
line, and. in all except Dinornis and
In Ehea, Struthio, and Dromceus the mouth of the
groove remains permanently open.
The Eustachian
aperture of each
is
conspicuous, lying on either side of the
In Apfenjx they
processes.
Irnsc
of the basipterygoid
In the
nearer the middle line and are inconspicuous.
basithe
very wide apart, opening immediately behind
lie
Tinamous these apertures are
ptervgoid processes.
The
paroccipital notch
and deep.
much
As in Rhea,
it
is
is
more
more
or less wide in
all
or less completely
save Apteryx,
bounded on
where
its
it is
narrow
outer side by a
the paroccipital process
compressed, forwardly directed lamina of bone from
(p. 172).
Tinamous,
Ehea, Dronums, Casuarius, Apteryx, and the
for
foramina
nerves, the condyloid
occurs the vagus foramen for the ninth and tenth
and the occipital condyle.
exit of the twelfth nerve lie behind it, between it
Within
this notch, in Struthio,
the
behind the notch and much nearer to
In the mnornithina' the vagus foramen occurs
cluster of apertures, that of tlie vagus
the condyloid foramina, forming, with them, a little
The carotid foramen in the I)inornWuda; lies
being much the longest of the group.
further
it occurs some distance
in front of the notch, in all the other forms
immediately
forwards.
fontanelles occur in many of these
Traces of both anterior and posterior basicranial
The former
the same skull.
PaliEOgnathine sku!ls, sometimes both are indicat.'d in
basipterygoid
line between the anterior ends of the
is the^small pit in the middle
2 c
XV.— PART V. No. % .—December 1900.
VOL.
,
ME. W.
174
processes
the latter
;
is
PYCRAFT ON THE MORPHOLOGY AND
P.
more often
slit-like
and
lies in
the middle of the basitemporal
platform.
The parasphenoidal rostrum, which
Apterycc, Casuarius, Bhea, Dromceus,
is
more
of great length,
and rod-like in
thin
is
or less inflated in the Dinornithidce, Struthio,
and Tinamous.
—
The Lateral Surface of the Cranium. The tympanic cavity is moderately deep
The mouth of the cavity
all save Aitteryx, in which it is comparatively shallow.
somewhat lozenge-shaped
in Dinornithidce, ^pyornis, and more or
less circulur in
in
is
the
other forms.
It is
bounded
posteriorly, in every instance,
by the base of the paroccipital processes,
save only in the Tinamous, where, on account of the greater relative smallness of the
the
process,
tympanum
stretched along
is
its
outer border.
free
externally by the free inferior border of the squamosal.
Tinamous,
In
In Ehea this process has shifted somewhat further iorwards, and
much
;
is
this recess in front,
the formation of
cavity to
taken by
continued forwards as a long,
this, is
form the anterior tympanic
it is
much
recess.
This
smaller in jTJpyornis.
wing of the parasphenoid bounds the aperture of
or alisphenoidal
and the pro-otic forms
its
is
posterior region of this cavity lodges the apertures
Bhea, Dromceus, and Casuarius;
largest in
The pre-temporal
The
the anterior, in front of
spacious, tunnel-like pneumatic
aperture
place
its
This second process may be regarded as a flange-like
shorter.
downgrowth of the squamosal.
of the internal ear
Rhea and the
save
all
bounded
edge of the zygomatic process.
this border represents the posterior free
a second but
It is
its roof,
posterior and ventral walls.
the basi- and exoccipitals share in
The basitemporal
plate and the
highly pneumatic basisphenoid receive the flnal termination of this recess beneath
the pituitary fossa, as
is
shown when the
skull
is
part in the formation of this cavity can of course only be
of very young nestlings.
Each
The bones taking
seen in section.
made out
recess curves gently forward to
meet
its
in
the skulls
fellow of the
opposite side.
In Apteryx this recess
is
exceedingly small, and appears to
There
not entirely, within the basisphenoid.
two
is
for the
lie,
most
part, if
no intercommunication between the
recesses.
The
roof of the tympanic cavity
surface of the quadrate.
Behind
formed
is
this
is
for the
most part by the large articular
a conspicuous cavity, the roof of which
is
perforated by numerous pneumatic apertures leading into the diploe Ixtween the prootic
and squamosal.
It
represents the superior tympanic cavity.
Mesiad of
this is
the fenestral recess containing the fenestra ovalis and rotunda and the aperture of
the posterior tympanic recess.
tympanic
In the larger Dinornithidce there
is
a small superior
recess.
The Squamosal Prominence.
the paroccijiital process.
—This
is
bounded
in front
by the temporal
It passes dorsad into the parietal,
and
is
fossa,
behind by
continued downwards
PHYLOGENT OF THE PAL.^OGNATH^ AND NEOGNATH.E.
to
form the zygomatic process.
is
of considerable length, and runs
In Casuarius, DromcBUS, u^pyornis, and Rhea this process
down
to
overhang
cup-shaped
to within a short distance of the
In Rhea
fossa for the articulation of the quadrato-jugal bar.
downwards
175
runs forwards and
it
this articulation.
In Dromanis, except D. ater, the anterior border of the squamosal prominence bears
a strong protuberance.
In Stndhio and Apteryx the zygomatic process
very short.
is
In Binornithidce the middle region of the squamosal
is
much
inflated,
making the
squamosal prominence very conspicuous.
The temporal
fossa
is
more
or less conspicuous in all the PaJceot/nafhce, ss.\e Apferyx.
In Casnorius and DronKBus
it
is
narrow and deep.
In the latter
its
limits
ai'e
sharply detined.
In the Dinornithido' the size of the fossa varies, but
general direction
In Rhea
it
is
generally wide and deep,
its
backwards.
wide and shallow, and with
is
is
resembles that of Rhea, but
is
In Cryptiiri
linguiform impression.
a
it
shallower.
In Strvthio narrow, moderately deep, and slopes obliquely backwaids.
^pyornis has the same type of
fossa as Stndhio, but deeper
and much more sharply
defined.
In Apteryx
far forwards.
it
is
It is
represented by a shallow but sharply defined depression extending
peculiar from the fact that
not bounded anteriorly by a post-
it is
orbital process.
The
post-orbital
process
in
Casiiarius
is
strongly developed
;
it
forms a broad
linguiform process extending downwards to within a short distance of the quadratojugal bar.
In Dromceus novce-hollandia' and D. ater
In Emeus, amongst the Dinornifhida',
In Rhea
it is
In Sfruthio
relatively small,
it is
it is
its
downward extent
is
much
less.
even larger than in Casiiarius.
somewhat pointed, and directed
slightly backwards.
but slightly developed, and runs forward to blend with a shelf-like
projection of the irontal overhanging the orbit.
In Cryptiiri
The
it is
triiiemiiial
represented only by a minute prickle.
foramen
is
a conspicuous aperture lying, in Casiiarius, at the bottom
of the temporal fossa at the point where the outer angles of the alisphenoid
d
pre-temporal wing meet, and in the same transverse plane as the basipterygoid processes.
is
Immediately behind
this, in
Drommis, and
less distinctly in
Apteryx and Rhea,
a smaller aperture leading into the tympanic cavity.
The
size.
orbits in Casuarius, Uroma^us,
and Rhea are
all
more or
less
alike in form and
All are strongly walled-in behind by overhanging and laterally projecting post-
orbital proci sses,
and are in front protected and increased by long backwardly directed
spurs from the lachrymal.
Struthio differs markedly from the above in that the spurs of the lachrymal over-
2c2
ME. W.
176
P.
PYCRAFT ON THE MORPHOLOGY AND
arching the orbits are short and blunt, but are connected with the frontals by a more
or less imperfect chain
of supra-orbital ossicles, as in
would seem, from the number of
skulls
which
A specimen
obliterate all trace of their independence.
however, has this chain on
one side
looking up the subject
I find it
to
in
it
fig.
have escaped the notice of
was
first
Museum,
The
3).
the Rothschild
developed (PL XLII.
pei'fectly
seems
existence of this supra-orbital chain
On
Generally
Crypturi.
tlie
have examined, that these supra-
with one another and the lachrymal and frontal so as to
orbitals fuse completely
writers.
I
recent
all
described by Cuvier [14] in 1799.
Hildebrand [37] in 1806 seems to have rediscovered them. Meckel [56] and other
writers at the beginning of the century were also aware of their existence.
There
is
a distinct, but small, pre-sphenoid fossa.
and raised high above the rostrum, as
^pyornis and Binornithidce agree
orbital
region of this
The
optic foramina are distinct
in JEpyornis.
in the
fused with the frontal to form one uniform plate of bone.
happen
filled
Struthio
in
the
if
The
form of the supra-orbital ledge.
pre-
formed externally by the lachrymal, which has completely
is
between
fossa
This
what would
just
is
backward spur and the nasal
its
were
up.
In yEpyornis,
as in Struthio, the optic
Below the
parasphenoidal rostrum.
foramen
is
lacerate fossa,
raised liigh above the level of the
and between
it
and the trigeminal
foramen, the surface of the combined alisphenoid and pre-temporal wing
inflated
and roughened by numerous elongated spine-like processes
much
is
attachment
for the
of muscles.
In Dinornithidw the optic foramina are deeply overhung by the outstanding orbitosphenoids
;
in this respect
resembling Aj)teryx.
In Apteryx the orbits are small and tubular, without post- or pre-orbital processes,
and the
interorbital region of the frontals I'educed to its smallest possible limit.
In Crypturi there
pre-orbital
is
no lacerate or pre-sphenoid fossa or post-orbital process
;
the
backward spurs of the lachrymal are wanting, and the interorbital region
of the frontal
is
deeply notched, being cut away to within a short distance of the
interorbital septum.
The
lacerate fossa in
however,
it is
some Divornithidce takes the form of a deep
represented only by a shallow depression.
pit
;
In and around this
generally,
lie
certain
of the cerebral nerve-apertures worthy of note.
In Lromceus and J^pyornis these are very distinct.
first
division of the v. (orbito-nasal)
and
slightly
and the
vi.
In the former those of the
(abducent) nerves
mesiad of the former, and both external to the
lie,
rest.
the latter below
That of the
iv.
(pathetic) lies mesiad of and slightly above the in., being divided only by a narrow bar
of bone from the
IV. it is
11.
(optic).
The
only separated from the
triangle with the in.
and
vi.
il.
III.
oculo-motor
lies directly
by a narrow bony bar.
foramina,
lie
below the
Lowest of
all,
iv.
;
like the
and forming a
the foramen for the arteria ophthahnica interna.
—
PHYLOGENY OF THE PAL.'EOGNATH.E AND NEOGNATH^.
In many Dinormthida' these apertures
lie closely
177
packed together down the mouth
of a deep lacerate fossa.
In most of the other Pald'ognntJm more or fewer of these foramina become more
or less confluent one with another and with the optic foramen.
—
The EtlimoiiJal Region. The mesethmoid, in all but Dinornithidce. and Apteri/x,
forms the thin median septum known as the interorbital septum. It is continued
forward into the pre-lachrymal fossa as far as the cartilaginous septum nasi.
Its
ventral border rests upon the parasphenoidal rostrum
its dorsal border affords support
;
to the frontal
and
ethmoidal plates.
as
This
nasals.
does by means of paired, lateral, horizontal, ecto-
form
the inferior aliethmoids, to
perforated superiorly in
communication
it
These, the sitpen'or aliethmoids, curve outwards and downwards,
tlie
the walls
of
chamber.
the olfactory
It
is
dried skull so as to place the two olfactory chambers in
but Apteryx, Casuarius, and Crypturi.
in all
In Dromwiis and Casuarius
the only region that
ossifies
whole of the outer wall of the aliethmoidal region
tlie
—
The lower
deeply invaginated.
is
region of this in-
vaginated wall forms a large, outstanding, antorbital plate, to the outer free edge of
which the lachrymal
is
closely applied
(PL XLIV.
lig.
The
1 a).
thus reduced to a narrow slit-like cavity, whose outer wall
is
olfactory
chamber
is
scroll-shaped, the scroll
being formed by the invagination before mentioned.
This invagination lodges the Harderian gland, and
fossa.
Opening above and behind
is
this is a small supra-orbital fenestra.
In Rliea the invagination of the aliethmoidal wall
Harderian
fossa,
though
passes gently
larger,
called therefore the Hardeiian
less
is
backward
sliarply
marked, and the
Except in
into the orbit.
very old specimens, the greater portion of the wall of this fossa remains unossifled.
The
supra-orbital fenestra,
when
ossification is complete, is large.
In Struthio only the superior aliethmoidal wall
region forms the antorbital plate; this
the roof of the olfactory chamber.
There
is
ossified.
The
postero-inferior
continued upwards and forwards to form
is
is
a supra-orbital fenestra.
In Crypturi the antorbital plate stands out at right angles to the mesethmoid as
a narrow bar of bone.
The upper portion
of Rliea in being deeply invaginated, so
the mesethmoid, thus forming a
chamber
In
There
aliethmoidal wall resembles that
so as to force
huge Harderian
to the smallest possible limits.
all
of the
much
is
fossa
it
inwards almost on
to
and reducing the olfactory
a well-marked supra-orbital fenestra.
the Paheognathce, save the Crypturi, the ossified ectoethmoidal plate extends
backwards, on either side of the dorsal border of the interorbital septum, as far as the
brain-case.
its
Thus only the lower region of
whole extent
and in Apteryx.
as in Neognathce.
septum
is
is
visible in the skeleton, not
most noticeable in the Dinornithidce
In these the olfactory chambers are of enormous
backwards nearly or quite as
upon the
this
This feature
far
as
size,
the optic foramina, thus so far
orbit as to obliterate the interorbital septum.
This
extending
encroaching
last is traceable only
MR. W.
178
P.
ON THE MORPHOLOGY AND
PTCKAI'T
in the Bhiornithidce as the vertical plate dividing the pre-sphenoid fossae.
There
is
a
well-marked supra-orbital fenestra.
The
Dinornithidce are apparently
peculiar in the development of a pair of well-
developed triangular processes, forming a horizontal plate projecting from the ventral
border of the mesethmoid, and extending outwai-ds to the level of the lachrymal.
much
In Apteryx the ossified ectoethmoidal plates form two oblong,
on either
side of the head, nearly filling
up the very small
inflated bullae
orbit, the horizontal
plates
the mesethmoid sweeping outwards, downwards, and inwards, the ventral edge
fusing with the mesethmoid immediately above the parasphenoidal rostrum. Forwards,
of
the part corresponding to the autorbital plate affords support to the lachrymal, as in
Bromceus.
As
"One
the late Prof. T. J. Parker has pointed out [71],
characteristics of the skull oi Apteryx
the mesethmoid
is
marked obliquely
one from another, and I propose
posterior turbinals.
:
in the olfactory
vertical folds, while a fifth
The
The
of the most striking
When
the extreme complexity of the turbinals.
removed there are seen
portion of the nasal cavity.
anterior turbinal
is
is
chamber proper four well-
continued into the narrow or respiratory
three hindmost of these folds are perfectly distinct
to call
them, respectively, the anterior, middle, and
fourth and fifth folds are intimately connected with the
I call the
uppermost of the two, which forms part of the olfactory
which extends forwards, and has no
region, the anterior accessory turhinal, and that
olfactory function, the ventral accessory turbinal.
" All the turbinals, with the exception of the ventral accessory, are covered by
Schneiderian membrane, and are therefore analogous to the ethmo-tnrbinals of a
The
ventral accessory turbinal
maxillo-turbinals of
"
is
is
covered with ordinary mucous membrane, and belongs
merely respiratory portion of the nasal chamber
to the
On
mammal.
;
it
may be compared with
the
mammals.
each side of the ventral edge of the mesethmoid, in the vomerine region,
a slender rod of cartilage, imbedded in connective tissue, and lying parallel to, and
either immediately dorsad or slightly laterad of the dorsal edge of the trough-like
vomer
it
can be readily
removing the vomer.
The
It is obviously the vestigial cartilage of Jacobson's
by
carefully
organ
."
.
.
ventral accessory turbinal does not appear to ossify in the adult Apteryx skull.
There are no
ossified turbinals in
The lachrymal
Its
prepared in a well-macerated skull
in
Casuarius
upper limb, when
is
the other Palceognathce.
crescentic in form, with convex border forwards.
visible, is very
overhanging ledge to the
orbit.
Its
broad and deeply notched.
lower limb
is
flattened
It
forms a broad
from before backwards and
extends downwards to within a short distance of the quadrato-jugal bar.
junction of the upper and lower limbs
passage of the lachrymal duct.
with the autorbital plate.
it
is
At
the
perforated by a large foramen for the
In the adult, the inferior inner lateral border
is
fused
179
PHTLOGENY OF THE PAL.EOGNATH.E AND KEOGNATH-E.
In
Bromaus
developed,
of the upper limb is s^'eatly
the outer fork of the >-shaped notch
supra-orbital process
a long, backwardly directed, spine-like
forming
notched below the lachrymal ioramen.
with the
this notch is short and confluent
In I) novwhlllandm the inner fork of
The outer fork is produced
the antorbital plate.
free ed-e of the ventral border of
In D. irroratns
quadrato-j ugal bar.
downwards far beyond this, and rests upon the
quite
of the prong are subequal, and do not
and B ater the outer and inner forks
continued
of the antorbital plate, which is
reach the level of the ventral border
it
to the maxiUo-palatal process, where
downwards by means of a slender column on
(PI
XLII
fi-
2).
The
inferior limb
is
expands.
,.
,
^-
•
i
outer fork of the upper hmb is comparatively
In Struthio (PI. XLII. fig. 3) the
furtherlaterally than in the above mentioned,
short and blunt, and projects more
of large bony scales l>ing between its
more it differs in that it is joined by a series
supra-
recalling the
and the supra-orbital ledge of the frontal,
Late in life the scales fuse with the laciuymal
orbital chain of bones in the Tinamou.
The
thus enclosing a large supra-orbital foramen.
in front and the frontal beliind,
whose
of a slender backwardly curved rod,
lower limb of the lachrymal takes the form
bar.
ugal
to join the inner border of the quadrato-j
free end is curved sharply outwards
ossiculum
is attached a larger
To the inner side of this outwardly directed free end
and the
witli the lachrymal on the one hand,
posterior extremity
palatinnvi, which in the adult fuses
antorbital plate
Below the
on the other.
ventral border of
its
inner end comes
m
contact with the palatine.
completely fused with the frontal
jEmjornis and Dinomis have the lachrymal so
The upper limb in both has
distinguishable.
and antorbital as to be with difficulty
The lower in Binormthda^ is much
completely merged into the supra-orbital ledge.
being
lachrymal foramen is imperfect, its outer bar
flattened antero-posteriorly.
wanting.
In some,
e. g.
The
Ammalopteryw,
it is
perfect.
Its ventral
bar.
truncate and articulates with the maxillo-jugal
enabled to examine,
In the only yEinjornis skulls I have been
been broken away.
border of the lachrymal
In both these groups the anterior
extremity
tliis
is
squarely
lower limb has
.
rests against a
marked, spine-like, maxillary nasal process.
by Parker [71], from young specimens,
In Apferyx the lachrymal is described
well-
as
"a
surface of
a shell-like central portion applied to the
small irre-ular bone consisting of
descending
ascending portion which articulates with the
the aliethmoid, and of an
In the
foramen."
lachrymal
It is perforated obliquely by the
the nasal.
process of
Its anterior border is often traceable
completely fused with the aliethmoid.
and the maxillary process of the nasals.
as a thin suture between it
indistinguishably fused with the nasal, and forms
In the Crypturi the lachrymal is
There are no
in front.
prominent outstanding process bounding the orbit
adult
it is
therewitli a
backwardly projecting spines.
The
ventral limb fuses with the antorbital plate.
ME. W.
180
The Cranial Cavity.
P.
—The
PYCKAFT ON THE MORPHOLOGY AND
metencephalic fossa in
save Dirwrnis and JEpyornis,
all,
takes the form of a moderately deep basin, the floor of which
The
very considerable extent.
Posteriorly
pituitary fossa.
it is
and
below the pro-otic
which
slightly posterior to,
to a
Its
pierced by the vagus foramen,
is
one or two condyloid foramina.
lie
Mesiad of the internal auditory meatus, and anterior to the same,
foramen
upwards
tilted
continued backwards on to the occipital condyle.
postero-lateral border, immediately
to the inner side of,
is
portion forms the posterior wall of the
anterior tilted
the abducens
is
(vi.).
In xSpyornis and Dinornis the floor of this fossa
The cerebellar fossa
otics bound it laterally,
is
almost
is
flat.
ventrally
passes into the metencephalic fossa.
it
beliind by the free edge of the occipital foramen, and in front by the
The
the tentunal ridge.
The
roofed by the parietal and supra-occipital bones.
late
Prof. Jefl'ery
It is
pro-
bounded
median portion of
Parker, in his masterly monograph, states
that in Dinornithidce the supra-occipital region of this fossa
Immediately above the
cerebellum."
grooves corresponding with the gyri of the
internal auditory meatus lies the floccular fossa.
The
appears to vary individually in Dinornithidce.
It
marked by " transverse
is
size of this,
according to Parker,
of considerable depth in the
is
other Pal(p,ognath(e and in Tinamous.
The
mesc7icephalic fossa
is
a deep basin-like depression in all save Dinornithidce and
Apteryo:.
It appears to
bounded
group.
It is
of the
tentorial
beyond the
be more sharply defined in Casuarius than in any other
ridge, behind
of this fossa,
mouth
by
the pro-otic.
(v.)
between
of this
it
in
forwards
the
considerably
middle
The
line.
leaves by a large aperture excavated out of the ventral border
it
and the
and
pro-otic,
of this foramen, below the rim of
orbito-nasal nerve
It extends
which bounds
level of the pituitary fossa
trigeminal foramen
member
above by a very prominent shelf of bone, forming the lateral portion
(v.'),
which has
its
its
exits in
is
of considerable
size.
Just within the
anterior border, lies the aperture of the
an almost obsolete lacerate
fossa.
In
Dromceus the trigeminal and orbito-nasal foramina are distinct. The tentorial ridge,
bounding the fossa superiorly, is almost as strongly developed as in Casuarius.
In Struthio this fossa
so shar|)ly
nerve
defined.
lie close
is
The
relatively smaller
and shallower, neither are
boundaries
its
apertures of the trigeminal foramen and the orbito-nasal
together.
somewhat more sharply defined than in Struthio,
but in none of these does that portion of the tentorial ridge bounding the fossa
superiorly form anything more than a low ridge.
The aperture for
In Dinornithidce this fossa is relatively ill-defined and shallow.
Ehea and Crypturi have the
fossa
the trigeminal foramen steals away a large portion of
nasal aperture opens as in Casuarius,
separating
it
from the pituitary
fossa.
is
its
tunnel-like, and
external wall.
bounds the
The
orbito-
fossa anteriorly,
—
PHTLOGENY OF THE PAL.^OGNATH.E AND NEOGNATH^.
In Aptery.v
oblong
in
seen at
its
it
appears to be reduced
shape
—
best.
to the
its
long axis liorizoutal,
It
lies
entirely
vanishing point.
— and
behind
more
extremely small,
or less sharply defined,
pituitary
the
of
the level
It is
181
pituitary fossa,
below by the rim of the metencephalic
The
projecting shelf-like ledge from the tentorial ridge.
its
postero-dorsal border, the orhito-nasal in
In A. haasti the outer wall of this fossa
common
to the trigeminal
The ]}ituitari/ fossa
is
its
fossa,
It
fossa.
bounded above by a low tentorial ridge, in front by a ridge of bone dividing
when
is
from the
it
and behind by a huge
trigeminal foramen opens at
anterior border.
almost entirely absorbed by a large aperture
and orbito-nasal nerves.
in Casuarius is a
deep pocket-shaped depressimi.
mouth
Its
is
behind, and the
thereof forms the dorsum sellce
Within
the mouth of this fossa [Linornis) or on its rim
pre-pituitary ridge in front.
(Drommis) is the oculo-motor (iii.) foramen. The pathetic (iv.) foramen lies above and
The rim
directed slightly forwards.
to the outer side of the oculo-motor, often
serves also for the passage of the internal
it
Anterior to the prepituitary ridge
ophthalmic artery.
in
the middle line
a narrow
is
which are the confluent o/jtic furariwui.
continued upwards vertically and again expands to tbrm the
knife-like optic platform, on either side of
The
optic platform
is
pre-optic ridge, which in
Dromwus, and
slightly less obviously so in Casuarius, joins
the tentorial ridge.
In Struthio
pituitary fossa
tlie
is
of great depth
;
the pre-pituitary ridge
and optic
platform are well marked.
In Dinornithidce the fossa
The
optic platform
is
is
comparatively shallow, but of considerable circumference.
very wide.
In Ajjteri/x the pre-pituitary ridge
is
The
but feebly developed.
optic platform
is
not distinguishable from the pre-pituitary ridge, which divides the optic foramina one
The pre-optic ridge is well developed and overhangs the fossa, the
from another.
mouth
of which, in consequence, has a semicircular outline.
here that the bottom of the fossa
spherical, depression,
In Tinamous
The
should be mentioned
produced forwards into a yet deeper, almost
overhung by the pre-pituitary ridge.
this fossa
deep and tubular
is
in form.
In Apteryx, and to a lesser extent in the Crypturi, the cerebellar contour
the cerebral from the remaining
The
fossae, arises
tentorial ridge,
from the
which serves
turns sharply upwards and forwards to meet
to cut
level of the pre-optic ridge
trends outwards and backwards to the level of the top of the pro-otic
;
from
Drommis
XV.— FART
V.
No.
is
ofl'
and
this;^point
the dorsum
selliE.
portion of the ridge which forms the upper boundary of the mesencephalic
Casuarius and
is
in the mid-dorsal line, over the region
corresponding with a vertical line passing upwards from
VOL.
all,
the cerebral lies entirely in front of the cerebellai-
scarcely distinguishable from the cerebral.
it
In
cerebral fossw are sharply defined in all the Pala-oguathce save Apteryx.
with the exception just mentioned,
fossa.
is
It
Ihat
fossa in
of great depth, and projects into the cranial cavity like a
lU.
December, 1900.
2 d
MK. W.
182
deep
This
shelf.
is
less
PTCRAFT ON THE MOEPHOLOGY AND
P.
marked
In Dinornis aud Cryptitri
in Ehea.
it
is
but feebly
developed.
The
cerebral fosste in Dinornithidce do not
infundibuliform olfactory
taper forwards to finally pass into the
on the contrary, are of great
fossae, but,
size
and width
in this
region, reducing the olfactory fossae to the dimensions of shallow pits.
The median bony
(/alii,
is
extremely
and only feebly
falx,
continued forwards from the tentorial ridge to the crisfa
well
developed
and Dromceus,
Casuariiis
in
There
Struthio and Dinornis and Tinamous.
in
less
Ehea,
so in
no trace of
is
it
in Ajjteryx.
In Castiarius aud Dromwiis the
fossae
bear distinct and deep impressions for the
temporal and frontal lobes, a low ridge indicating an indistinct Sylvian
The
olfactory fossce in
save Dinornis and
all,
A^iteryx, are paired,
separated one from another
tubular, infundibuliform chambers,
fissure.
by
more
or less
a crista galli.
Anteriorly they are closed by a more or less perforated plate for the passage of the
olfactory nerves.
In Dinornis the olfactory
end of the large cerebral
fossae
take the form of shallow pits lying at the anterior
The
fossa.
cribriform plate
is
large.
backwards into the
In A^iterjix these fossae are of great size, and pass insensibly
cerebral fossae.
They
are not, in the dried skull, shut off anteriorly from the turbinals
by a cribriform plate as in other Eatitce, but continued forwards directly into the
olfactory chamber.
The Premaiilla.
The premaxilla
in
all
the Falceognatlice
median, and not paired as in
all
is
peculiar, in that the
other birds
(PI.
XLII.
nasal
In
1-3).
figs.
process
all
is
save
Struthio there are well-developed palatine proceses.
In Casuarius the beak
is
more or
less
laterally
The body
compressed.
—that
portion of the premaxilla from which the nasal, maxillary, and palatine processes are
given off
—
is
short,
and more or
less
deeply grooved on either
side, so
that the distal
end of the nasal process seems to be continued forwards into a strongly marked ridge.
The maxillary processes extend backwards as a pair of slender splints to the
The palatine processes are a pair of short
the lachrymal on either side.
level of
narrow-
laminae extending backwards to a point corresponding with a vertical line passing
upwards behind the top of the
free
They enclose
end of the median nasal process.
anteriorly a small chink-like palatine notch.
In C. australis there
is
a small pre-
narial septum, corresponding to that described by Parker in Dinornis, as the " strong
ascending keel, formed posteriorly of paired plates, but solid in front and gradually
diminishing in height towards the top of the beak."
This keel
is
derived from the
dorsal surface of the palatine processes.
The
nasal process
in the inflated,
is
rod-like, its free
end passes backwards into a tunnel-shape
mesethmoid mass forming the characteristic casque of these
birds.
PHTLOGENT OF THE PL^OGNATH^ AND
In
Dromaus
the beak
than in Casuarius, and, as
continued forwards to the
divide
it
The
genus,
this
in
tip of the
end
rests
The maxillary
anterior region of
The
relatively longer than in Casuarius.
is
lateral grooves
be
which
nasals.
is
received
The
tip
of
processes are splint-like, and extend backwards to the level of the
tlie
orbit
—beyond
body of the lachrymal.
tlic
closed caudad, as in
The
far as the level of the anterior
In Casuarius,
it
sliould be mentioned,
it
falls
In iJinornis the median nasal process
The maxillary
more widely separated one
chink-like anterior palatine
Casuarius, by the vomer.
backwards as
ventrally.
it
upon the mesethmoid.
from another in the middle line than in Casuarixs.
is
Posteriorly
median borders of the
palatine processes are short pointed lamellae, rather
foramen
larger
from the maxillary processes.
nasal process
free
is
process seems to
the rod-like nasal
beak by reason of paired
into a shallow groove excavated out of the
its
18o
'
The premaxiUary body
flattened dorso-ventrally.
is
NEOGNATIIJ^:.
is
This process extends
border of the posterior narial a-perture.
short of this.
markedly flattened and expanded dorso-
processes are very short, not extending backwards beyond
the level of the hinder end of the antorlifal fossa.
The palatine 2^rocesses take
the form of flattened laminae, extending backwards, as in
The
JJromceus, to the level of the anterior end of the posterior nares.
alveolar borders
are deeply pitted.
Tlie anterior palatine
foramen
is
small and cordiform.
It is closed
In Emeus
parasphenoid rostrum, and posteriorly by the vomer.
it is
above by the
more completely
closed than in any other PalaognatlKc by a pair of processes from the postero-internal
angles of the palatine processes, which meet in the middle line, serving at the same
time as articular surfaces for the anterior ends of the probably paired vomers.
The prenarial septum
border
is
is
most
clearly
marked
bilaminate, but the laminae rapidly fuse
in JJinoriiis.
to
Its
postero-ventral
form the single median vertical
septmn.
In .Mpyornis the nasal process rises sharply from the body of the bone, giving the
beak a peculiar conical form.
extensions
The
of the palatine and maxillary
Museum skull or that
Museum skull there is a
the r>ritish
British
alveolar border
is
deeply pitted.
processes cannot
The backward
be made out in either
Hon. Walter Eothschild. In the
prenarial septum, but the laminae composing this
Itelonging to the
have not fused to form a median plate as in Dinornis. Instead, as indicated, they
remain widely separated, and enclose between them a long tunnel-like cavity extending
forwards to the tip of the beak.
In Rhea the premaxiUary body
much
depressed dorso-ventrally.
is
of greater length than in any other Ratite, and
It is
grooved dorsally on either side of the distal end
of the nasal process, as in the other forms.
The
nasal process
is
very broad and flattened, but tapers
more
or less
caudad
2d
'J
ME. W.
184
The
maxillary,
process,
The
it
at,
confluent for about half of
is
and terminates
palatine process
terminating
In this
which
styliform,
is
PTCKAPT ON THE MOEPHOLOGY AND
P.
of great
is
length with the palatine
in the anterior region of the antorbital fossa.
size,
extending beyond the maxillary process, and
or falling but little short of, the anterior border of the posterior nares.
Dromwus.
agrees with
StnitJiro differs
from
consequent enormous
all
the other Ratitce in the absence of a palatine process, and the
size of the palatine
The whole premaxilla has
a
body of the bone
is
so that the
its
foramen.
unique triradiate form
;
the radii are of great length,
reduced to the smallest possible dimensions.
The nasal process is broad and flattened, tapering somewhat abruptly posteriorly.
The maxillary processes are long and styliform, terminating at the level of
the
anterior region of the orbit.
In
free
Crijpfirri
the form of the premaxilla
end of the palatine process
precisely similar to that of Bhea.
is
brought
is
posterior nares, and interlocks with the distal end of the palatine.
it
differs
from Bhea.
Apteryx
is
the
The
palatine foramen, as in Rhea,
most abnormal of
all
is
In this last point
long and narrow.
The whole
the PakeofinatJue.
produced forwards into an elongated probe, curved slightly downwards,
which the external nostrils
The
to the level of the anterior border of the
beak
is
in the tip of
lie.
In the adult skull the fusion of the separate elements, not only of this region of the
beak alone, but of the skull generally,
is
make out their
The body of
to its
so
complete,
that
it
is
impossible
to
boundaries.
the premaxilla
is
reduced
smallest possible limits, being com-
posed only by the confluent end of the rod-like nasal process, and of the combined
maxillary and palatine processes, which for this reason are but delicate rods.
The
nasal process takes
running backwards
to the
the form of a long slender rod, grooved ventrally and
mesethmoid.
Its posterior region rests
upon the elongated
nasal bones.
The
length,
maxillary and palatine processes are fused throughout the greater part of their
and form elongated flattened
representing the palatine foramen.
rods, divided in the
Posteriorly there
palatine spur, representing the palatine process.
middle
is
line
by a narrow groove
given off a short splint-like
The extremest
limit of this
falls
considerably short of the anterior end of the antorbital fossa.
The maxillary process can be traced backwards splint-wise as far as the level of the
foot of the lachrymal.
Thus the maxillary processes are of about equal length.
nasal,
The
and nasal process of the premaxilla above, and the maxillary process of
the premaxilla below, serve to form a long narrow groove leading forwards into the
anterior nasal fossa, this last having shifted forwards to the extreme end of the beak.
This nasal furrow bears a strong resemblance to that of the Ibises, the difference being
that the nasal fossa lies behind instead of in front of the groove,
seems
as in Apteryx.
It
to suggest that the shifting forward of the nasal aperture has followed the course
—
PHTLOGENY OF THE PAL.EOG.NATILE A\D XEOGNATH.E.
•of
the orbito-nasal nerve, accompauying
it
185
bring the sense of smell
to its tip, so as to
to aid the sense of tcnich found in the Ibises.
It is interesting to note that in the very early
the position of the external nares
seems
to
near the middle of the beak.
is
Its
forward shifting
be involved in the growth of the elongating beak.
TJie
The maxilla
in
line so as to
MaxiUo-jvgal Arch.
takes
Casaariiis
interim- half has the outer
tlie
form of an elongated flattened
and inner borders turned upwards
mouth of
of the
this
is
meet
to
form a long thin-walled tunnel tapering forwards
The rim
antrum.
embryo, as shown in Parker's figures,
splint.
in the mid-dorsal
to a point.
This
is
The
viewed ventrally.
the posterior region of the
which
in the ventral
floor of this tunnel,
maxilla.
is
the
produced backwards into two short spikes;
the outer forms a strong dorso-lateral and the inner a lateral spike, conspicuous
skull
Its
viewed ventrally,
This wider region
— the
is
when
the
twice as wide as
the antrum,
floor of
view presents a deeply hollowed free posterior i)order forming the
anterior boundary of the posterior narial aperture,
is
the maxillo-j_)alatine process.
The whole inner border of this process is grooved for the reception of the vomer.
The maxilla extends forwards above the maxillary and palatine processes of the premaxilla.
The latter process is received into a shallow depression in the maxillowhich bone can be seen lying between the palatine and
The backward extension of the maxilla is considerable. It runs
palatine, a raised ridge of
maxillary process.
below the quadrato-jugal bar on the outer side and the palatine on
its
inner side to a
point slightly beyond the level of the free border of the hinder end of the posterior
nares, or, to take another standard, to within a short distance of the base of the vomer.
The quadrato-jugal fossa
the
is
name which
I
have applied
to
indicate the large
space enclosed by the quadrato-jugal bar on the outside, the palatine and pterygoid in
the inside, and the hinder end of the maxilla in
In Casuarius
it
fi'ont.
does not extend forwards beyond the hinder end of the posterior
narial aperture.
In Uroiiueus the maxilla
extend so far backwards.
is
relatively
The
Casuarius.
Its dorsal wall,
l)orders of the maxillo-palatine processes rest
The quadrato-jugal
in Casnarins,
and does not
maxillo-palatine processes are wider but very
shorter antero-posteriorly than in
reduced in length and capacity.
much wider than
The antrum,
moreover,
is
in
consequence,
fenestrated.
is
The
much
much
mesial
upon the vomer.
fossa extends forwards to within a short distance of the
middle
of the posterior narial aperture.
In Dinornithidw the maxillo-palatine region of the maxilla
anterior end
maxilla.
is
wedged
Ventrally
it
in
between the maxillary and
appears as an
obhmg
is
Its
|)alatine processes of the pre-
flattened plate.
border bounds the anterior end of the posterior nares.
short and broad.
Its
Its
postero-iuternal
posterior border
is
notched or hollowed and forms the anterior boundary of the quadrato-jugal
cither
fossa.
ME. W.
186
From
its
PTCEAFT ON THE MORPHOLOGY AND
P.
dorsal surface in Dinornis there arises a large hollow shell of bone, the inner
wall of which embraces the triangular plate of the rostrum.
shell
The
cavity of this
— the antrum — opens posteriorly by a large foramen into the quadrato-jugal
It articulates
with the palatine by a short recurrent process from
bony
fossa.
postero-internal
its
angle.
In Emeus there
is
AVhat corresponds
no antrum.
to its inner wall in Dinornis exists
as a flat concavo-convex vertical plate applied to the anterior border of the
mesethmoid
triangular process and the rostrum.
In Rhea the maxilla
Viewed
with
dorsally
its
it is
sharply
is
almost entirely represented by the maxillo-palatine process.
seen to take the form of a
flat,
end running forwards
pointed
triangular, perforate plate of bone,
to within
a short distance of the
anterior end of the external narial aperture.
This more pointed region rests upon the
bounded externally by the maxillary process of the premaxilla.
Its postero-mesial border is bounded by the vomer.
Its hinder end presents free
postero-internal and external borders.
The former lies more or less markedly forward
palatine process, and
than the
latter,
is
and bounds the posterior nares
anteriorly.
The
latter lies
more
or less
considerably candad, and bounds the anterior end of the quadrato-jugal fossa.
postero-internal angle
is
produced
backwards
into
a long spiue,
which
is
Its
closely
applied to the outer border of the palatine, extending backwards for a considerable
distance.
Its
postero-external border
underlies the quadrato-jugal bar.
is
From
also
produced backwards spine-wise, and
the dorsal surface of the maxillo-palatine
plate there arises a long slender style which, directed
upwards and backwards, ultimately
articulates with the anterior angle of the lachrymal,
lachrymo-nasal
birds this
is
(antorbital)
and thus serves
from the nasal fossa (PL XLIII.
done by the descending maxillary process of
tlie
fig.
The
—
^just
divide the
In
9).
all
other
The antrum
nasal.
represented only by a minute aperture at the base of the vertical rod
nasal process of the maxillo-palatine
to
is
— the lachrymo-
described.
quadrato-jugal fossa extends forwards nearly as far as the level of the anterior
end of the posterior nares.
In Struthio the maxilla
processes, the inner of
running
ventrally
splint-shaped, produced
is
which runs dorsad over the
below
the
quadrato-jugal
nearly as far as the body of the premaxilla.
and hatchet-shaped
for their reception.
The
;
their mesial borders
There
is
posteriorly into two spine-like
distal
bar.
The
end of the palatine, the outer
Anteriorly
clamp the vomer, which
a small shallow antrum, the roof of
anterior border of the maxillo-palatine
extends forwards
it
maxillo-palatine processes are small
is
grooved laterally
which
is
fenestrated.
process bounds the enormous palatine
foramen posteriorly.
The
ossified style, described in
Rhea
as dividing
tlie
lachrymo-nasal from the nasal
here represented by a ligament springing from the external dorsal angle of the
mouth of the antrum and inserted into the short descending process of the nasal.
fossa, is
PHTLOGENY OF THE PAL.^OGNATH.E AXD NEOGNATILE.
The
187
quadrato-jiigal fossa extends forwards to withiu a short distance of the anterior
border of the posterior nares.
In Cryptiiri the maxilla resembles that of Rhea, but
much narrower and
is
bounded
is
There
unfeuestrated.
is
its
maxillo-palatine process
The lachrymo-nasal
no antrum.
anteriorly by a descending process of the nasal
is
fossa
and not by a spur from the
maxillo-palatine as in Shea.
In Ajjtcryx the maxilla
extending forwards
to th(^
is
of great
size, its
maxillo-palatine processes, seen vcntrally,
middle of the elongated beak, and backwards along the outer
horder of the palatine to within a short distance of the level of the free end of the
They
orbital process of the quadrate.
forwards, only by the narrow vomer.
from another iu the middle
are separated one
Posteriorly there
The proximal
articulation with the quadrato-jugal bar.
There
with the external border of the short palatine.
line,
an external lateral spur for
is
mesial border
is
no antrum.
Immediately in
is
closely fused
front of the distal end of the palatine the maxillo-palatine process occasionally sends
inwards a very
process
short
articulate
to
with
the
vomer,
which
apparently
much larger vomerine process found in Struthio.
The ([uadrato-jugal is so reduced in size in Casuarius as to be represented by little
more than a nodule of bone, scarcely projecting beyond the limits of the glenoid cup
corresponds with the
of the quadrate.
The
jiKjal in
Casuarms
portion,
and projects above
spine.
It
is
connected by a close vertical suture with the quadrate
it
at this part
in the
form of a more or
prominent
less
extends forwards as far as the middle of the lachrymo-nasal fossa.
In Dromwus, seen from the outside, the quadrato-jugal scarcely exceeds that of
Casuarius
(PI.
XLIII.
On
4«).
fig.
the jugal for about one-third of
its
the inner side, however,
length.
the rim of the quadrate cup on the inside.
same
it
extends forwards along
does not extend beyond
The forward extent
of jugal
is
about the
as in Casuarius.
Rhea and Strutkio resemble Dronueus
In Apteryx the quadr<(fo-ju<jal
middle of the zygomatic arch.
along
In Casuarius
it
the
dorsal
is
in the size
and relations of these two bones.
a long slender style, reaching as far forwards as the
The jugal
is
long and slender, extending forwards
aspect of the quadrato-jugal as far as the level of the anterior
extremity of the palatine, which corresponds with a line drawn transversely across the
vomer about
its
middle.
In Crypturi the quadrato-jugal
is
as
much reduced
as in Casuarius.
The
mesial
border of the proximal end of the jugal articulates, as in this form, with the quadrate.
The Vomer, Pterygoid, and Palatine (see also
In Casuarius the vomer
C. c. sclateri,
and
C.
c.
is
p.
2u6).
(PI.
of great length and slenderness.
beccarii
it is cleft
XLII.
In
6".
tigs,
c.
u-8.)
salmdorii,
from behind forwards for more than half
its
ME. W.
188
PYCEAFT ON THE MOEPHOLOGY AND
P.
length into two slender rods grooved dorsally
divided
:
is so-
the cleft not extending more than half the distance of the posterior nares, in
other forms
tile
in C. hennetti only about one-fourth
;
both types, the
continued forwards beyond this point.
it is
cleft of
terminating in
the
C. c. australis differs
from
the vomer being rather less than half the total length and
region
end of the posterior narial aperture.
of the anterior
Anteriorly the vomer becomes laminate and wedged in between the maxillo-palatine
and passing forwards
processes,
The
upon the palatine processes of the premaxilla.
rests
palatine foramen by the great forward extension of the
In a ventral view
vomer appear
—before
wedged
to be
vomer
is
almost obliterated.
complete anchylosis has taken place
feet of the
Removal of
between the pterygoid and palatine.
in
— the
latter, or a dorsal view, however, reveals the true relations of these parts.
be seen that the right and
It will
this^
then
limbs are closely ajjplied to the external ventral border
left
of the twisted pterygoid, extending backwards to a point very near
its
The
middle.
pterygoid can be traced forwards along the mesial border of the vomer as a slender
On
spike for some considerable distance.
a dorsal view, the right and
left
grooved
limbs of the vomer are well displayed, and show that each has been twisted outwards
so that its inner free border
comes
The
look doisally, and the latter ventrally.
to
flattened surface of this twisted region
is
closely applied to the ventral surface of the
vomer and pterygoid form one common rod
The dorsal border of the twisted end of the vomer is
anterior end of the pterygoid, so that
articulating with the quadrate.
slightly thickened and, traced
of the pterygoid.
From
backwards, trends downwards beneath the outer border
the vomer backwards, the sutural surface
is
continued as a
shallow groove along the outer ventral border of the pterygoid, terminating at the
level
the
of
basipterygoid
processes.
It
pterygoid that gives the impression that the vomer
bones,
when
the skull
In Dromceus
one-third of
(PI.
is
is
wedged
in
of palatine
and
between these two
viewed from this surface.
XLII.
its total
union
ventral
this
is
7) the
fig.
length.
"feet" of the vomer extend forwards
Anteriorly
it
for
about
expands into a broad oval plate tapering
The
again forwards to terminate, as in Casuarius, near the body of the premaxilla.
end
processes
maxillo-palatine process overlaps
the
underlie
with the pterygoids and palatines posteriorly are
ventrally.
it
Its relations
distal
dorsally
precisely similar to those Avhich obtain in Casuarius.
remarked, the palate of Dromceus
consequently represents, as
all
we
is
As
;
the palatine
a matter of fact,
it
should be
even simpler in type than in Casuarius, and
shall see, the simplest
arrangement of these bones of
the Palaw/nathce.
After these, the simplest palate amongst the remaining Palaeognathine forms
is
that
of Struthio.
The vomer
it
of Struthio
(PI.
XLII.
fig.
8)
is
a degenerate
does not extend beyond the middle of the enormous palatine foramen
it is cleft,
but has entirely lost
all
Anteriorly
structure.
;
posteriorly
connection with the pterygoid, faUing short thereot
PHYLOGENV
THE
Ol'
PAL.EOGXATir.T-:
by some very considerable space.
AND
has assumed a trough-like form, the hollow of
It
On
the trough closely embracing the parasphenoidal rostrum.
is closely
embraced by the maxillo-palatine processes, whicli
A
facets.
but
similar,
perfect,
less
189
XEOaXATJI.E.
arrangement of
tit
vomer
either side the
into specially
these
parts
roughened
obtains
the
in
Cassowary.
we should have a
of Casuarias and Dronwux than any
vomer were produced backwards on
If the
type of palate more nearly resembling that
pterygoid
the
other Ratite.
In Rhea the vomer
cleft posteriorly for
is
rather less than half
and each terminating
length, and
its
duced anteriorly into two large blade-like laminfB, widely separated
in the
is
median
These anterior lamime are embraced
in a pointed extremity.
The
between the palatine processes of the premaxilla and the maxillo-palatines.
rostrum
parasphenoidal
appears
in
median
the
line,
terminating at the level of their free anterior ends.
Struthio the anterior end of the rostrum
is
In
between
far
beyond the rostrum,
takes the form of a pair of vertical laminae, fused anteriorly,
rostrum, upon which
it
slides;
processes,
Casuarins, Bromceus, and
and of the vomer terminate at the same point.
free end of this
these
entirely concealed by the vomer.
two Ibrmer the free end of the vomer projects
proline,
the forward anterior horns
In the
in the latter the
The body of the vomer
and moulded on to the
lie in
a horizontal plane;
the feet are obliquely truncated, their external lateral borders suturating with the
pterygoid (PI. XLII.
The
fig.
>).
vomer
true relations of the
a study of the dorsal view
;
to the
Casuarius or iJromau.s, the foot of
This
pterygoid.
last
is
pterygoid cannot be
the vomer underlying the anterior end of the
B/iea rod-shaped
in
form, but
in
dorso-ventrally compressed and closely applied to the
in adults
it
seems to become
made out except from
they will then be seen to be precisely what obtains in
less
vomer
the
sutural
at least in
;
compressed and yet more pointed.
surface
young
is
birds,
In a ventral view
of the skull the vomer would appear to suturate with the palatine only and not the
pterygoid, the former being apparently thrust in between them.
Examined from the
dorsal surface, however,
it
exact form of which will be discussed presently
either side, and
is
will be seen that the palatine
—underlies
— the
the foot of the vomer on
continued backwards along the pterygoid in the form of a long
pointed spur, concealing
all
but the proximal end of the pterygoid, so that, seen
ventrally, as already remarked, the
vomer and pterygoid would seem
to be shut off on(>
from the other.
This
is
a fact of considerable significance, inasmuch as this isolation of the vomer
a point wherein
gnathce.
the Neognathd' appear to
But, as I have elsewhere shown,
(p. 206), this is
only an apparent
In Cryjjturi the
VOL. XV.
all
— PART
v.
vomer
is
when
and
is
differ
from the Paheo-
discussing the
palatine of Ehe((
agree,
dift'erence.
precisely similar in form and proportions to that oi Rhea.
No. 1\.— December, 1900.
2
li
MR. W.
190
Furthermore,
The
but
it
PYCEAPT ON THE MOEPHOLOGT AND
P.
agrees with the same in the nature of
by
palatine, in the part played
little
from that of Rhea.
narrower from side to
with regard to the pterygoid and vomer,
it
the fact that
Its chief difference consists in
and that
side,
rehitions with the pterygoid.
its
terminates in a pair of spines,
it
The form
inner, instead of an outer pterygoidal only as in Bhea.
differs
much
it is
an outer and an
of the palatine, as
a whole, will be described latter.
In Linornithiche,
Emeus
e. g.
(PI.
XLIII.
the vomer takes the form of a
fig. 3),
embracing the
parasphenoidal rostrum.
pair
of perfectly distinct vertical laminae
They
are continued forwards only for a very short distance beyond the
anterior end of the posterior narial aperture.
greater vertical
the paired
extent of
from that of Bhea
It differs
laminae,
(2)
level
are
these
that
in
of the
(1) in the
often
free
throughout, and (3) in that there are no anterior horns running forward between the
In Emem, the palatine
palatine processes of the premaxilla and the maxillo-palatine.
processes send inwards a pair of flattened spurs to embrace the sides of the anterior
end of the vomerine
plate.
Posteriorly, the feet of the
The
relations
vomer suturate with the pterygoid exactly
between these bones and the palatine
Emeus
inturned under-flooring region of the palatine in
than in Bhea, projecting backwards to the level of
In Apteryx the vomer
half
its
length.
relatively short,
is
and
its
cleft
Indeed, the
more strongly developed
pterygoidal spur.
form and relation are described on
in
Casttarius
is,
is
the same.
from behind forwards
Its
The fterygoid
also
is
as in Bhea.
p.
205.
seen ventrally, an elongated, crescentic rod of bone
dorso-ventrally compressed, with convex surface directed towards, and partly
and
on, the parasphenoidal rostrum,
the palatine.
the vomer
Anteriorly,
— which
is
it
its
wedged
terminating thereon in a sharp spine.
below
concavity, for the most
Its
in
between the pterygoid and palatine
with
— and
proximal end articulates with the quadrate,
mesiad and anterior to this articulation
;
resting
part, articulating
runs forwards along the inner side of the hinder limb of
at this point
the orbital process
about
for
is
Seen dorsally, the pterygoid
articulation with the basipterygoid process.
a second for
is
fusiform,
not crescentic, a large median tube overlapping that part of the palatine which on a
ventral view appears within a crescentic hollow.
cannot be seen without
overlapping
the removal
of the
Thus the
palatine,
true shape of the pterygoid
which
discloses the dorsal
lip.
In Dromceus (PL XLII.
fig.
7) the
Its relative length is aboTit tlie
anterior region of
its
form and relations of the pterygoid are simpler.
same, and
external lateral border
The
it
terminates anteriorly in a point.
is
directed upwards in the form of a thin
vertical plate affording a sutural surface for the external vertical plate of the vomer.
The
Its
posterior region of this border
internal, mesial
border
is
is
straight
and
affords attachment to the palatine.
convex, the convexity facing the angle between the
PHYLOGENY OF THE PAL^OGNATH^ AND NEOGNATH.E.
191
parasphenoidal rostnim and the basipterygoid processes.
The palatine is but loosely
attached to this bone.
Tlie free end of the vomer undcrfloors its anterior end.
In Stncthw the form of
Dromams: the
It lacks
pterygoid (PI. XLII.
tlic
wherein
points
differs
it
8)
resembles that of
closely
points wherein
are
the npstanding vertical plate from
fig.
it
also degenerate.
is
antero-external latend border, and
its
it
has fused with the palatine.
In lihea the pterygoid (PI. XLII.
in size,
fig.
and now takes the form of a
facets for articulation with the quadrate
twisted around
laterally
own
its
parasphenoidal rostrum.
axis,
Yentrally
it
8) has
undergone a very considerable reduction
The sliort upper limb bears
and basipterygoid processes.
The longer is
-shaped rod.
s
and continued
is
grooved.
anteriorly, to the superior border of the vomer,
forwards
In the Linornithidce,
comes
reaches the
it
This grooved surface
and posteriorly
the postero-external angle of the posterior extremity of the
posterior end of the palatine
till
to the
is
applied,
upper surface of
Thus the
palatine.
to underlie the posterior extremity of the
vomer.
Emeus, the form of the pterygoid very closely resembles
The points wherein it differs are such as must be regarded as repre-
that of Rhea.
e. g.
senting a more primitive condition: the chief of these lies in the greater width dorsoventrally
;
the pterygoid of
Its relations
Emeus
possessing a broad ledge along
its
mesial border.
with the vomer and palatine are practically the same as in Ilhea.
In Cnjidtm the jjterygoid
is
rod-like as in llliea, relatively longer,
much
straighter,
and more slender.
In Aptenjx the pterygoid
upon and
fuses with
p.
elongate, laminate, and pointed in front where
the vomer.
palatine, so that the precise
the adults (see
is
204).
It
form of the pterygoid
Proximally,
it
rests
anchyloses very early and completely with the
it is
tightly
as a
whole cannot be made out
wedged
in
in
between the basipterygoid
one side and the orbital process of the quadrate on the other, the
pterygoid having the appearance of being " morticed
in between these two jioints
process on
tlie
"'
of contact.
')l\\&
palatine
'\Vl
Casuarius takes the form of a roughly triangular plate.
articulates with the pterygoid
the maxilla.
Its connection
and vomer, and the anterior half of
with the pterygoid
is
its
Its
by means of an overlapping suture,
The
the pterygoid underlying a broad shelf of bone given off from the palatine.
vomerine suture
is
that of a simple apposition
junction with the maxilla.
base
outer side with
of contiguous edges, as also
is
the
In the adult the palatine and pterygoid become indistin-
guishably fused proximally.
In Dromceus the palatine resembles in form that of Casiiarius.
fenestrated.
the pterygoid,
Its suturations
'lliere is
In Bhea the palatines
differ
only in that there
is
XLII.
fig. ti)
is,
however,
no overlapping shelf above
no fusion between pterygoid and palatine as
(PI.
It
in Cnsuarius.
are broad thin laminte, roughly quadrangular
2e2
MK. W.
l'.J2
PYCEAFT ON THE MOEPHOLOGY AND
P.
with the external horder produced backward into a pointed spur which runs
in form,
along the pterygoid, and forwards for a considerable distance along the maxilla.
mesial border
is
cut away anteriorly so as to embrace the external border of the
instead of underlying
it,
as
done by the remaining
is
posterior portion.
Its
Its
vomer
form
is
seen best from the ventral aspect of the skull.
In the DinorvUhida/,
but by a
g.
e.
Emeus, the form of the palatine resembles that of Bhea,
come
tension the body of the bone has
little
to
be vertical instead of hori-
zontal in position, thus giving a rod-shaped superficial appearance.
In the Cryptiiri the main body of the palatine
anteriorly produced external
border
lateral
is
as in Rhea, horizontal, but the
is,
much
of
greater length, and takes the
form of an elongated and slender rod suturating anteriorly with the maxilla.
In Struthio the palatine (PL XLII.
the main body of the bone
peculiar, however, in that
external lateral border
rod, but
is
is
is
many Carinatw
twisted into the form of an almost vertical scroll.
it is
fused at
its
in that
It is
The
base with the overlapping pterygoid.
produced forwards, as in Tinamous, into an elongated slender
which fuses anteriorly with the base of the hastate maxillo-palatine process.
In Apteryxthe palatine (PL XLII.
rod,
8) resembles that of
fig.
diff"ering
in
relations
its
from
5) takes the
fig.
form of a more or
forms.
the other associated
all
The
fused with the maxillo-palatine process.
greater part of
its
less flattened
Anteriorly,
external border
it
is
anchylosed with the maxilla.
The remaining
the pterygoid.
Dromceus
What
corresponds with the posterior region of the mesial border in
here free, and runs in a backwardly transversely oblique direction across
is
the skull.
bone anchyloses with the external border of
posterior end of this
For the precise relations of the bone however, see
The Quadrate.
(PL XLIII.
figs.
204.
p.
10-12.)
The quadrate in Casuarius has its various processes very sharply defined.
The otic process is very slender, tapering gradually downwards into the body of the
At its base, where it merges into the main body, is a large pneumatic foramen,
bone.
The head for articulation with the pro-otic and
it opens on the posterior surface.
squamosal
The
to
is
single,
oblong in form, with
its
long axis directed backwards and inwards.
inner and outer mandibular condyles join mesially, the two projecting backwards
form a conspicuous spur.
The
difference in
tlie
level of the
two condyles
is
but
slight.
The
orbital
superior border.
process
At
its
is
well
defined,
moderately large
extreme base, and on
its
inner side,
and
is
free,
with a sinuous
a small demi-facet for
the pterygoid.
The outer
surface of the quadrate
is
very closely applied to the long stout zygomatic
process of the squamosal, which terminates only a short distance above the level of the
cup-shaped quadrato-jugal
fossa.
PHYLOGENY OF THE PAL.EOGA'ATJLE AND NEOGNATH^.
The quadrate
Lromwus
ot
193
but slightly from that of Casuarim.
differs
The outer
and iuner mandibukir condyles are
on distinctly different levels, tlie inner being
ventrad of the outer, shorter from witliout inwards and wider from before backwards.
The backwardly
projecting condylar spur
less
is
marked and formed
entirely by the
outer condyle.
The
orbital process
is less
In the JJinoruithnhv,
hollowed above.
Emeus, the quadrate resembles that of Dromanis in the
between tlie outer and inner mandibular condyles and in the form of
difference of level
the orbital process.
e. g.
This process
is,
however, peculiar in that
requistioned for the articulation of the pterygoid.
peculiar in that the middle region of
the outer condyle looks more directly
The pneumatic foramen
sometimes present
is
The inner mandibular condyle
articular surface
its
whole inner surface
its
downwards than
is
in Droiiiojus.
A posterior foramen
mesiad, and not posterior.
however,
is,
in the Uinorn/'fhidce.
they closely resemble those of Dronums.
other forms so far described, in that
there projects from
The
is
produced downwards, whilst
In Strut/do the inner and outer mandibular condyles are widely separated
pterygoid.
is
The
its distal
;
otherwise
orbital process differs from that of the
end turns sharply upwards, and
in
that
inner ventral border a sharply defined articular facet for the
its
otic process is relatively longer
mesial and posterior foramina are
present,
than
in the other forms.
though
the
Both the
may apparently be
latter
sometimes absent.
In jEpyornis the mandibular condyles resemble in form and disposition those of
some respects
Stridhio, and in
from
differed
all
other genera in
is
it is
taken was kindly lent
me by
is
Dronurm
;
its
process
orbital
of
is
lies
articulation with the quadrate.
In
Crjjijturi
the form of the quadrate
this
facets.
to
otic
process
is
The external
The inner is
is
vertical
very short.
the long axis of the skull.
is
cpiite
it
and
Its
The
The
does not project beyond the level
peculiar in the development of a
The
bent backwards at a sharp angle, and forms with the orbital process a
Y-shaped bone.
is
rather closely.
cingulum projecting backwards above the external mandibular condyle.
otic process is
but
articulated with the pterygoid
immediately below the articular condyle.
very short, blunt, and very wide,
its
distinct
The
long axis at right angles
posterior pneumatic foramen
have
missing, but from
of these two last the outer
looks forwards and outwards, the inner ventral.
head has
is
to
The specimen from which
mesial.
broken up into two distinct articular
is
similarly subdivided but not so distinctly
articular
it
end
seems
Dr. Forsyth Major.
In Bhea the quadrate resembles that of
mandibular condyle
orbital process
Its free
probable that
The pneumatic foramen
Sfrufhio.
description
great breadth.
its
the sharp edge of the proximal end
as in
The form of the
Diiiornis.
The
more slender
;
external form of the orbital process resembles that of Stnithio
the
method of
articulation with the pterygoid closely resembles
ME. W.
194
P.
PYCRAFT ON THE MORPHOLOGY AND
The form of the t)iUer and inner mandibular condyles is not unlike
The inner condyle is, however, more nearly spherical tlian in any of
that of Binornis.
that of Linornis.
the other forms herein described.
is
Apteryx possesses a readily distinguishable quadrate. The height of its vertical axis
less than that of a line passing backwards through the orbital process, which is very
long.
The
mandible present the same ground type as
articular condyles for tlie
and Casuarius, but the external condyle presents a tendency
to split into
B/iea,
In
it
should be mentioned, there
all
peculiar in that the pterygoidal articulation
more than half of
for the pterygoid
Arising at
than in the other Pakeognatluv.
it
is
to the orbital process for
In Apteryx the articular surface
the
is
its
base of the internal mandibular
extends outwards on to the inferior border of the base of the orbital process,
otic process
elongated,
its
line,
is
Its external
fits.
Superiorly transversely
has two fairly distinct articular facets.
mesial extremity
with the pro-otic.
middle
length.
rather more sharply defined
taking the form of a deep hollow into which the pterygoid closely
The
facets,
process
a considerable space between these two points.
the Pala'0(j)iath(e the quadrate
extends upwards on
condyle,
is
two
In Struthio and
closely applied to the outer side of the quadrate.
is
Dromceus
The zygomatic
the region nearest the quadralo-jugal bar being scooped out.
of the squamosal
in
expanded into an almost circular head and articulates
extremity runs obliquely
and affords an articular surface
down
to the shaft towards the
for the alisphenoid
and squamosal.
The pneumatic posterior aperture is sometimes absent.
The separation into pro-otic and squamosal heads in the other Pakeo(jnatlm
is
only
very feebly indicated.
The
exoccipital does not contribute towards the articular surface for the quadrate in
Apteryx.
(PL XLIII.
The Mandible.
The mandible
is
expanded
in Casuarius is long, slender,
to afford
fig. 9.)
and sigmoidally curved.
an articular surface for the quadrate.
This
is
Posteriorly,
it
moderately deep,
and with sloping sides meeting at a sharp angle at the bottom of the fossa. Internally,
The surface for the external condyle
it is produced into a blunt internal anyular p>TOcess.
forms a long narrow facet running parallel with the external border of the cup that
;
for the inner
forms a band-shaped
facet,
hollow from within outwards.
without the inner end of this internal facet
angular process
is
is
a pneumatic foramen.
Immediately
The
posterior
only feebly developed.
In Dromceus the internal and external angular processes are larger than in Casuarius.
The rami are almost
straight,
and the symphysis
jaw agrees with that of Casuarius.
is
very broad.
In Pinornithidce,
e. g.
In other respects the
Linornis, the articular
PIITLOGENY OF THE PAL.EOGNATIL^ AND NEOGNATH^.
surface for the external condyle of the quadrate
The
cup-shai)ed.
is
19-')
internal and
external angular processes are feebly developed.
In Bliea the form and disposition of the articular surfaces agree with those of
The
Droma'us.
ramus bears a
internal angular process
in
The symphysis
large.
very broad.
is
Tlie
small, perforate posterior lateral vacuity.
Strutliio in the
however,
is
form of
its
mandible
closely resembles
ratlier
greater relative slenderness, in that
its
Dronwus.
It differs,
si^moidally curved, in the
it is
greater relative size of the expanded posterior portion, and in
tlie
elongated form of
the articular facet for the external condyle of the quadrate.
The rami
jEpyornis also resembles Droma'us.
however, more laterally com-
ai'e,
pressed, have an elongated anterior lateral vacuity,
broad dentary border, and the
a
articular facet for the internal condyle of the quadrate situated at the
cup.
Tlie internal angular process
bottom of a deep
well developed and bears a large pneumatic
is
foramen.
In Crypturi the mandible resembles that oi Bl/ea.
The
posterior angular process
is,
however, longer in Tinamous, and the posterior lateral vacuity appears to be wanting.
Apteryx
is
differs
from
all
the other forms in the great length of the symphysis, which
about half the length of the ramus.
The
in Stnithio,
it is
reduced
to a
mere
is
well developed.
small and rod-shaped.
Sometimes, as
Tlie internal angular process
coracoid in all the Palteognathine
forms
vestige.
is
The dentary suture
is
always distinct.
The Ilyoid.
Except
in Apteryx, only the cera to-branchial s
appear
to ossify in the Palseognathine
Epibranchials and the true hyoid elements appear to remain
forms.
cartilaginous.
permanently
In Apteryx the basihyal appears to ossify at least occasionally.
The Skull of the Nestling.
The Cartilage-bones.
The youngest of the Palseognathine skulls in my possession is that of an embryo
The cartilage-bones are in all the stages herein described completely
Droma'us.
ossified.
The
basi-occipital, in the
young Lrommis mentioned above,
from the neighbouring bones
It is
bounded
(PI.
antero-laterally,
XLIII.
and
fig.
in front,
it
forms only the ventral part of
this.
perfectly distinguishable
5).
by the basi-temporal plate of the para-
sphenoid, and postero-laterally by the exoccipital.
the median portion of the occipital condyle.
is
It
is
produced caudad
It is interesting to note,
It is entirely
shut
off
from
to
form
however, that
the occipital
foramen
196
Mlt.
W.
PYCKAFT ON THE MORPHOLOGY AND
P.
by the exoccipital portions of
Seen
dorsallj',
Anteriorly
it
which meet
this condyle,
the basi-occipital
above
line
it.
cut off from the pro-otic by a broad patch of cartilage.
is
only just touches the basisphenoid.
In the youngest Cassowary and Rhea skulls in the
of the basi-occipital have almost closed.
It takes,
Museum
collection, the sutures
however, a slightly larger share in
the formation of the occipital condyle, inasmuch as
formation of
median
in the
takes a small share in the
it
free border.
its
In a half-grown Aj)teryx australis mantelH
agreed very closely with
it
that
of
Drotmens, and, as in this form, the exoccipitals meet in the middle line above the basioccipital.
The exoccipital, externally,
(inner)
The share which
is
border
Its dorso-lateral
hollowed
to
form the
lateral
meeting
left plates
is
bounded by the
in the
in
and squamosal
Bhea
—
considerable
is
in Bromwiis,
:
in
Casii-
Bhea, extending
this last, in
The lower portion of the
to join the supra-occipital.
4).
middle line behind the basi-occipital.
parietal
and Apteryx, and by the squamosal only
backwards
boundary of the
posterior
its
The lower
foramen magnum.
(fig.
takes in the formation of the occipital condyle
it
Bromceiia the right and
is free,
portion of
border bounds the infero-lateral border of the epiotic
region of this border
ariiis,
The upper
relatively large.
is
antero-lateral border
and in the shape of a large paroccipital process bounds the tympanic recess
posteriorly.
angle
Its internal dorsal
lies
within the tympanic cavity, and helps to form
the inferior pro-otic articulation for the quadrate.
Internally, the exoccipital
The
is
almost entirely concealed by the opis- and pro-otics.
and Rhea, runs upward and forward
sitpra-occipitaJ, in Di'oma'us
blunt cone
in
;
and Apteryx
it
Apteryx the superior border
That portion of
comes into contact with the exoccipital
be
is
its
easily distinguishable,
its
;
;
in
Bhea
runs up-
it
outer (inferior lateral) border which
In the Penguin
really belongs to the epiotic.
but in the forms now under description
made out on a reference to the interior of the skull.
The epiotic is not indicated externally in the peculiarly
Penguins
In Dromwirs, Casuarivs,
gently hollowed.
cut off from the squamosal by the exoccipital
is
wards to join the squamosal.
this last
is
form of a
in the
distinct
can only
it
manner seen
in the
even in the skull of the youngest of the embryo Apteryxes in our collection
only indication
is
the supra-occipital.
the venous canal which points the boundary line between this and
Internally,
it
is
fairly
skulls of this collection save those of the
In
distinct in all save Apteryx.
embryo Apteryx and Dromwifs,
it
all
the
has fused
with the pro-otic.
The
(PI.
pro-otic,
XLIII.
fig.
even in
the youngest
5) skulls, is
concealed by the squamosal.
in
which a large portion of
half-grown (82).
Ajiteryx (PI. XLIII.
fig.
3)
and Lroirueus
not visible externally above the tympanic cavity, being
In this
this
it
differs
markedly from the young Penguin,
bone remains uncovered
until
the bird
is
quite
—
THYLOGENY OF THE
The
I'AL.EOflN'ATHJE
AND NEOGNATH.E.
197
pro-otic completely shuts out the squamosal from the interior of the skull in all
In this type a large, roughly triangular plate appears immediately above
but Apterya'.
wedged
the pro-otic and
The
floccular fossa
in
between
tlie
parietal
deep, and oblong in
is
vertical to tiiat of the skull.
In Aptery.v
it
and alisphenoid.
but Apteryx, the long axis running
all
represented by a small but deep and
is
perfectly circular pit.
more
Externally, the pro-otic appears only within the tympanic cavity and affords two
or less distinctly separated articular surfaces for the quadrate
above and in front of the other.
articular surface furnished by the alisphenoid
to
Externally, articular surface for the quadrate
Thus
are
two
pro-otic,
one
a second
lies
be most distinctly traceable in
CasuariKS.
thei-e
the one lying slightly
caudad of the inferior
;
These appear
by the exnccipital.
facet furnished
:
Distad of the superior glenoid surface lies a similar
is
completed by the squamosal
one alisphenoid, and one squamosal
exoccipital,
articular facet for articulation of the quadrate.
The
otic
opistliotic
and
seen as a small oblong nodule of bone wedged in between the pro-
is
the exoccipital, and
bounded above by the
inferior
end of the
It
epiotic.
is
not visible externally.
The foramen for the vagus lies at its ventral extremity and between it and the exoccipital.
The hasis])]ienokl has in every case fused completely with the luiderlyingbasi-temporal
plate,
and
is
therefore
traceable
only from the superior aspect of the skull.
It
is
moderately thick in section, the body of the bone being thickened by a mass of diploe.
bounded behind by the
It is
Anteriorly, in Ehea,
it is
basi-occipital, laterally by the pro-otic
and alisphenoid.
produced forward into a thin vertical plate of bone,
cartilaginous presphenoid
;
in the other
forms this vertical plate
is
much
serves to form the inner half of the circumference of the foramen for
nerve
(v.),
outer
the
portion being
contributed
anterior region
is
scooped the pituitary
either side near
its
postero-lateral angle.
It
by
the
It
the orbito-nasal
Out
alisphenoid.
The abducent foramen
fossa.
to join the
shorter.
of
its
it
on
pierces
forms the floor of the anterior region of
the metencephalic fossa.
In Apteryx
expanded
The
it is
almost quadrangular.
laterally to
is
narrower behind than in
it is
front.
bounded by the parietal and squamosal behind, the orbital plate of
above, and the combined basisphenoid and alisphenoid as rings of the
alisphenoid
the frontal
In Casuarim and Uroiiiaus, and in Ehea,
form a pair of wings, and
is
parasphenoid.
In iJronueus, Casuaritis, and B/iea
it
bears a share in the formation of the jiost-orbital
process.
The
alisphenoid lodges the greatei- part of the mesencephalic fossa, and
contributes
a share towards the formation of the anterior wall of the cerebral fossa.
The orhitosphenoid
The presphenoid is
VOL XV.
— PART
V.
is
represented only by cartilage and
is
much reduced
in size.
represented by a thin vertical plate of cartilage resting upon
No. 12.
December, 1900.
2
i'
th(>
198
Mil.
W.
P.
PYCKAPT ON THE MOKPHOLOGT AND
parasphenoidal rostrum and wedged in between mesethmoid in front and the anterior,
compressed
laterally
plate of
vertical
the basisphenoid just described.
extends
It
upward and backward to the orbital plate of the frontal. From this region it sends
outwards and backwards a pair of cartilaginous wings, which represent the orbitosphenoids.
The mesethmoid,
a conspicuous interorhital septum.
?
Ostrich,
and
common with
in all but Apteryx, in
in the
the presphenoid region, forms
In the nestling Droiiuens, Casuaritis, Rhea, and
adult Tinamou, this
is
pierced by a large interorhital fenestra.
This fenestra appears to be excavated partly out of the large, backwardly directed
plate which unites with the orbital plate of the frontal, and partly (caudad) out of the
anterior border of the presphenoid.
In the nestling, the antero-ventral border
is
more
or less linguiform, and terminates at the level of the anterior border of the lachrymal.
From
this, there
runs forward a narrow, triangular, cartilaginous septum
nasi.
In the
Drommis, Binornithidce and Tinamous, ossification extends forwards
almost to the end of the rostrum, and terminates in a vertical truncate border, beyond
which there is a small cartilaginous septum nasi. In Rhea and Struthio the ossification
adult Casuarius,
,
extends quite to the free end of the rostrum,
relatively further forwards,
e.
/.
In these, the
terminates in a sharp point instead of a vertical truncated border.
cartilaginous septum nasi
The
is
wanting or greatly reduced.
and
dorsal border of the mesethmoid, in Drovneiis, Casuarius,
of considerable thickness and very slightly convex
olfactory fossa, extending outwards on either side
(PI.
XLIV.
;
it
Di)iornithid(e,
is
forms a complete floor to the
to the
In this particular, as will be seen
fig. 4).
and
orbital
later, it
plate of the frontal
differs
conspicuously
from Apteryx, wherein this region forms a thin median partition between two extensive
At the
cavities.
it
anterior end of the olfactory fossa, both in Uromceus and Casuarius,
sends upwards, and then backwards, a strong median pillar deeply grooved on either
side for the passage of the olfactory nerves.
described in Apteryx by
craiiii
of the lower vertebrates.
entirely
by
this
At the apex
tegmen
This corresponds to the tegmen cranii
Parker, and referred by Kitchen Parker to the tegmen
T. J.
ci-anii,
Thus the
crista galli, in these
the " tegminal process
"
ecto-ethmoidal plates
Drommis
— the aliethmoids,
as Jeft'ery
is
furnished
of Jeffery Parker.
of this tegminal process, as has been well
mesethmoid reaches the outer surface of the skull; here
two forms,
it
described by Parker,
tlie
throws out lateral horizontal
Parker proposed
to
call
them.
In
the upper surface of these plates forms a lozenge-shaped area, bounded by
the frontals behind, the nasals laterally, and the nasal process of the premaxilla in
front.
The
free edges of these right
downwards and inwards (PL XLIV.
and
left
aliethmoidal plates in the nestlings turn
become further extended
and intricately folded to form the lower region of the aliethmoid and the antorbital
In the nestling DrorncBus it should have been remarked that the median vertical
plates.
portion of thb mesethmoid is pierced by a large vacuity, the cranio-facial fissure.
fig. 4),
and
in the adult
PIITLOGENY OF THE PAL.EOGXATH.E AND NEOGNATILE.
Casuarhis
199
conspicuously from Lroinwus and the otlier forms associated here-
differs
with in that, that portion of the mesethmoid which in Lromceus, for instance, reaches
tlie
outer surface of the skull to form the lozenge-shaped plate already described,
surmounted by an independently
casque of the adult
— the
ossifying-
mass forming the centre of the characteristic
casque being furnished by the inflation
lateral portions of the
oi the nasals and that portion of
tlie
the nasal and median masses (PI.
XLIV.
is
frontal
figs,
bone which comes into juxtaposition with
o
&
4 a-h).
The median
portion often
extends so as to cover the parietal.
this casque is seen to
In the adult
and protected
tissue of great delicacy,
and development of
be built up of a homogeneous mass of cancellated
tliis
The form
in the living bird by a horny sheath.
casque varies considerably, and
is
one of the
factors
employed
in the determination of species.
The exact homology
of this casque has yet to be
made
That the
out.
lateral
portions are simply inflations of the nasal and frontal bones there can be no doubt, but
the origin of
median portion
tlie
structure superimposed
is
not so certain.
appears to be an independent
It
upon the mesethmoid, rather than an outgrowth of that element.
XLIV.
Stages in the growth and composition of this casque are shown in PI.
figs,
o
and 4 a-b.
In Rhea and the Tinamous, as in Dromceus and Casnarias, the dorsal border
mesethmoid comes
relationship
into chjsc relation
even
is
formed only by the tcgminal process.
Dromieiis and Casnarias
much
In
like,
But the passage
tliis
galli
is
for the olfactory nerves is
an exceedingly narrow space lying on either side of
restricted to
the
with the orbital plate of the frontal, but the
border being narrower, so that the crista
this
closer,
cjf
this process.
passage, though formed in precisely the
same way,
In
is
larger.
Ajjteri/x
and
the dorsal border of the
slightly concave.
It
is,
it
The tegminal
exceedingly narrow and knife-
is
moreover, relatively
than in Casnarias or even iJromaas.
the embryo,
mesethmoid
much
process
shorter antero-posteriorly
is less
appears on the surface of the skull as in Uronueas,
well-developed.
and
also, as
In
in this
form, sends out horizontal plates which form a lozenge-shaped shield between the frontal
and nasal bones.
The quadrate
Later in
in the
life all
trace of this shield
embryo of Dronueas
differs in
is
obliterated.
shape from that of the adult in
the lack of an ossified orbital process, and in that the external
is
mandibular condyle
<-siiapcd instead of transversely oblique.
In Apteryx
that
of the embryo appears to differ
in
nowise
from
that
of the
adult.
The
other skulls in the jNluseum collection are too advanced to afford any intermediate
characters.
The
the
articalar
is
a pyramidal nodule of bone,
base of which conceals
its
anterior
face;
bounded cephalad by
externally (in Rhea)
the coronoid
by the supra-
2f2
MK. W.
200
angular.
It furnishes
P.
PTCEAFT OX THE JNEOliPHOLOGY AND
In Dromanis and Apteryx,
the articular surface for the quadrate.
both angular and supra-augular take part in the protection of
The
stajjes is
its
external face.
the bony base.
represented in the dried skull only by
It does not
The
apparently offer any characters of value ironi a systematic point of view.
extra-
columella has been already admirably described by GadoAv, Parker, and others.
TJie Membraiie-hoiie><.
']^h.e
parietal, in Casuarius,
Dromceus, and IStruthio,
of bone, the anterior and outer borders of which
but sharp point which
is
wedged
in
ai"e
a transversely oblong plate
is
produced forward into a small
between that portion of the frontal which over-
hangs the post-orbital process and the process
itself.
form the base of the post-orbital process, the
main body of which
This parietal spur thus helps to
is
furnished by the
alisphenoid.
In Bhea and
and
muamous
fails entirely to
external angle
is
this antero- external parietal angle is not
reach the postorbital process (PL
XLIII.
produced forwards,
fig.
postero-
Its
9).
overlapped by an upstanding process of the squamosal.
Internally the parietal carries on the tentorial ridge from the alisphenoid upwards to
its
junction with the falx.
Behind
this ridge it is gently
roof of the cerebellar fossa, in front
scooped out
also hollowed out to
it is
complete the
to
form the posterior wall
of the cerebral fossa.
In Apteryx, as in the other forms, the parietal
border
it
is
bounded
entirely by the squamosal.
extends forward beyond
Internally,
it is
is
In
Its external
irregularly four-sided.
all
the other forms,
it
be noted,
will
this.
found to lack the tentorial ridge.
Its
The former
pass insensibly the one into the other.
hinder and external borders
the supra-occipital and a
skirts
portion of the pro-otic beyond, the latter rises gently from the hinder region of the
superior border of the pro-otic
to skirt the
—Avhere the hinder border may be said
into the formation of the inner wall of the
alisphenoid.
The
In the other forms,
e. g.
skull.
It
—upwards
entirely cut off
is
from the
Rhea, Dronuens, the superior borders of the
and the alisphenoid form the boundaries ventrad of
pro-otic
to cease
upper border of the small triangular area of the squamosal wliich enters
this bone.
frontal extends backwards so as to form the greater part of the roof of the
cerebral fossa.
Its
hinder border
is
mor-e or less sinuously curved,
and traced from
within outwards will be found, in Casuarius and Bromwus, to run transversely from the
middle line to the base of the post-orbital process, from which, however,
cut
oft'
by a very slender spur from the
external angle of the parietal,
upon the base of the
itself to
it
parietal.
it
is
actually
In Hhea, on reaching the antero-
turns abruptly upwards and outwards, so as to rest
post-orbital process itself.
Externally, the frontal
is
bent in upon
form a deep orbital plate which articulates caudad with the alisphenoid and
cephalad with the mesethmoid, eventually terminating in a long spike, in Rhea, resting
upon the mesethmoid, and overlapped by a greatly elongated
posterior extension of the
PHTLOGENY OF THE PAL^OGNATH^ AND
^'EOGNATH.t:.
201
Dromwus
the relations
nasals and the free end of the inner spur of the hichryniul.
are similar, but the uasal spur
is
In Casuariufi the forward extension of the frontal
Avith the
mesethmoidal region of the
The
antorbital plate.
lu
shorter.
orbital
greater part of
its
curtailed, since
is
plate, at the level
mesial border
of
it
origiu of the
tlie
much thickened and
is
being involved in the formation of the casque, and reiiresenting
this region
In some species
postero-lateral boundary.
terminates
this inflation of the
its
inflated
;
hindmost
bone extends backwards
as far as the parietals.
Fiu'thermore,
fact that the i'rontals
significant to note the
it is
fail to
meet
in
the
middle line anteiiorly, where they under-floor the median ossification of the casque
(PL XLIV.
Thus
fig. 4).
part in the formation of the roof of the cerebral fossa.
though
to take
It
would seem, from
this, as
gradually engrafting itself more and more
secondary ossification was
this
comes
a portion of the ventral border of the casque
much
completely upon the skull, by absorjition of the underlying bone in
as
bone has replaced cartilage in other cases
The extreme
so as to
way
in the cranial skeleton.
anterior end of the mesial border of the frontal
form the external
the same
is
gently hollowed out
boundary of the superficial lozenge-shaped plate of
lateral
the mesethmoid.
In
Ajjterya: the frontal
presents one or two points of difi'erence from that of the
remaining forms associated herewith.
border
Its mesial
bounded by the
joinis
parietal,
Immediately
aliethmoid.
fellow of the opposite side,
its
as usual.
Its
immediately behind the
line,
of the frontal gives
Cephalad of
fontanelle
is
oft'
of
tlie
bar
—between
frontal,
which
its
is
crista galli.
frontal sends
its
downwards and inwards
a
fellow of the opposite side in the
From
the outside, this orbital plate
a short slender plate to the aliethmoid.
plate
this orbital
bounded
hinder border
its
postero-veutral or lateral border joins the
in front of this, the
narrow plate of bone which, eventually, meets
middle
and
anteriorly
at its
is
This
a dee[) notch, the supra-orbital fontanelle.
by a narrow bar of bone, called the descending proces.'i
ventral extremity joins the aliethmoid.
upper and the outer wall of the aliethmoid
— the
Mesiad of
this
orbito-nasal nerve
passes from the orbit to the nasal cavity.
The squamosal,
nil
the forms herein dealt with, entirely conceals the pro- and
In the embryo
opisthotics.
quadrangular,
downwards
in
its
IJroDUi'iis,
antero-ventral angle
which forms the subject of
is
thickened on
is
as to render
immovable.
it
its
almost
produced into a blunt process which extends
to the level of the quadrato-jugal articulation.
the squamosal
this paper, it is
This downward process of
inner surface and so closely applied to the quadrate
In the embryo skulls of Dromceus, the squamosal and the parietal are connected
by means of a close suture, the juxtaposition of the two elements forming a perfectly
level surface.
Posteriorly the squamosal rests U2)on the outer border of the exoccipital.
and later fuses therewith.
Similarly the squamosal, anteriorly, rests U2)on the
ali-
202
]ME.
W.
p.
PYCEAFT ON THE MOEPHOLOGT AND
sphenoid, anchylosis therewith taking place
Thus
later.
it is
that the pio-otic comes to
be entirely concealed.
In
all ihis,
skull of this
Dromceus and Casuarius seem,
be
emarginate and
by
free,
than the Penguin.
less primitive
group recently described and figured by
to the dried skull simply
dee2)iy
to
me [82]
the squamosal was attached
articulation with the parietal.
its
exposing the pro-otic.
In a
hinder border was
Its
border was also
Its anterior
free,
not extending to the alisphenoid, but exposing between itself and this last a small
portion also of the pro-otic.
There
is
one great point of difference, however, between the squamosal of Bromceus
and that of the Penguin.
downwards
this spine
Except
In
the former,
into a large " zygomatic process," to
is
in
replaced by two small spines, as in
embrace the quadrate
all
Apteryx the squamosal takes no part
angle
antero-ventral
its
;
produced
is
in the latter,
the Neognathce.
in the formation of the inner wall of
the skull.
In
Apteryx the squamosal has
entered into a slightly closer relation wdth the
alisphenoid and exoccipital than in the forms described above, resembling in this
Thus the anterior border
respect the Neognathce.
convex border of the alisphenoid
;
is
concave and
fits
the corresponding
whilst the posterior border, though separated by
a narrow line of cartilage from the exoccipital, yet gives signs of effecting a close union
therewith immediately.
The squamosal appears on
the inner surface of the skull as a small triangular plate
above the pro-otic.
The nasal
in
Lromceus
especially so in the adult,
is
holorhinal.
where
it
The maxillary
exists only as a vestige.
process
It
is
is
greatly reduced,
produced caudad into
a long slender process resting on the frontal, and projecting beyond the lozenge-shaped
plate of the mesethmoid, to
process
is
which
binds externally.
Its
premaxillary or internal
of considerable length, extending forwards almost the whole length of the
nasal process of the premaxilla,
Dinornis
A
it
is
which protects them from above.
holorhinal.
skull of Megalajyteryx, in the collection of the
Hon. Walter Eothschild,
retains
form of the nasal
in the
sufficient of the traces of sutures in this region to indicate the
Linornithida^.
In
this skull,
it is
seen to be relatively
much
shorter than in Lromceus.
Its
extension does not pass the level of the horizontal mesethmoid plate and
pointed as in Bromceus.
plate projects
beyond
its
Its
maxillary process
wanting.
The
is
not so
ossified aliethmoidal
lateral border.
In Casuarius the nasal
is
schizorhinal
.
premaxillary process long, as in Bromceus.
is
is
backward
greatly inflated and takes
The maxillary
process
is
It is peculiar in that the
vestigial,
and the
body of the bone
part in the formation of the casque as already described
PHYLOdEXY OF THE PAL.EOUNATILE AXD XEOGXATH.E.
203
In Bhea the nasal must be regarded as liolorhinal, though the maxillary process
Inasmuch
absent.
the anterior nares,
as this process,
when
]n"esent, serves as the
boundary, caudad, of
place must be regarded as being filled by a vertical spike of bone
its
arising from the maxillo-palatine process to the anterior border of the lachrymal.
Struthio this vertical
nasal fossa
—
is
is
spike— in Rhea separating the
represented by a ligament which
In
anterior nares from the laclirymo-
attached dorsad to the free end of
is
the maxillary process of the nasal.
In Struthio the nasal
is
XLII.
holorhinal (PI.
fig.
Its
.3).
maxillary process
is
downwards on the maxillo-palatine by ligament.
as holorhinal, though the nasal cleft has extended
well developed, and continued
The Tinamida' may be regarded
backwards
The
The
so far as to lie level with the free
maxillary process
nasal
cleft
is
is
Above,
perfectly free.
is
it
it
bony
representing the
rod,
therefrom in being rod-like, very slender,
articulates,
by means of a short inturned process, with
the vestigial maxillary nasal process, whilst
articulates with the extreme postero-external border of the maxillo-palatine
process where
it
joins with the rod-shaped portion of the maxilla.
was regarded by Kitchen Parker
in all the skulls I
The
vertical,
It differs
the anterior border of the lachrymal and
ventrally
in-ocess of the premaxilla.
not wanting as in Rhea, thougli reduced to a mere vestige.
bounded externally by a
maxillary spike found in Rliea.
and
end of the nasal
This vertical rod
as the maxillary process of the nasal.
have examined, quite distinct from
It
is,
however,
tliis.
lachrymal, in Droniceus, sends backwards a very long, curved, rod-lilce process
to overhang,
and thereby greatly increase the
size of the orbit (PI.
XLII.
fig.
2).
It
sends downwards a large antero-posteriorly flattened limb, the outer border of wliich
is
produced
still
further
by a large foramen
downwards
The body
the quadrato-jugal bar.
into a short rod-like spine
which articulates with
of this ventral process of the lachrymal
for tlie passage of the
lachrymal duct.
is
perforated
The inner border
of this
ventral process articulates with the antorbital plate.
In
Casuarms the laclirymal
shortness of
its
differs
from that of iJroma'ns in the greater relative
external orbital process, which
is
scarcely longer than the internal.
A
further point of diff"erence lies in the fact that the external orbital process passes
insensibly forwards and
downwards
into a laterally compressed plate,
which, after
forming the boundary caudad of the posterior narial aperture, turns sharply backwards,
and becomes the antero-posteriorly compressed ventral process bounding the lachrymonasal fossa as described in Dro)nceus.
is
of
From
this
it
differs in that the
laminated portion
continued downwards to the level of the quadrato-jugal bar, instead of stopping short
this,
and sending downwards thereto a spine from
the lachrymal duct
is
smaller than in Broinceiis.
its
outer border.
Internally
it
The foramen
for
articulates witli the
antorbital plate.
In Rhea the lachrymal resembles that of Bromceus.
large.
The
lachrynio-nasal process
is
The
supra-orbital process
also antero-posteriorly compressed.
is
The foramen
ME. W.
204
for the laclirymal dvict
P.
PTCEAFT OX THE MOEPHOLOGT AND
Aery large.
is
quadrato-jugal bar and
is
This lachrymo-nasal process stops short of the
subquadrate ossiculum
connected therewith by a large
Internally the last eveut'ially fuses with
lachrymo-palatinum.
antorbital plate.
tlie
Its supra-orhital process is,
In Struthio the lachrymal resembles that of Bhea.
In two nearly adult skulls (one of which is
however, relatively shorter and thicker.
from the collection of the Hon. Walter Rothschild) this orbital process supports one
end of a chain of irregularly shaped ossicles which
Normally
frontal as far as the post-orbital process.
is
continued backwards along the
this chain of ossicles is so
completely
fused as to be indistinguishable from the frontals on the one hand, and the lachrymal
There
on the other.
a large supra-orbital fenestra bounded in front by the lachrymal
is
and behind by the frontal
The lachrymo-nasal
externally by a portion of this chain of supra-orbitals.
;
process
is
continued downwards as a sigmoidally curved rod to
On
terminate on the inner side of the quadrato-jugal bar.
extremity of the lachrymal there
lies
the inner side of this
a subcylindrical ossiculum lachrymo-palatinum
The lachrymal
articulating mesially, in very old specimens, with the antorbital plate.
foramen
is
represented only by a deep notch.
In Aptcryx the lachrymal
plate of
bone
is vestigial,
and takes the form of a small roughly triangular
closely applied to the outer surface of the aliethmoid
It is perforated
process of the nasal.
and the maxillary
by the lachrymal foramen.
The 2)re7naxilla. maxilla, jugal, quadrato-jugal, vomer, palatine, and pterygoid differ
from that of the adults already described
in no important respects
(p. 1S7).
The vomer, pterygoid, and palatine of Apteryx demand, however, a
too completely fused with
tion, as these in the adult are
render
it
make out
possible to
The pterygoid
is
the neighbouring bones to
their boundaries.
cephalad, being divided into two moderately long, pointed and
is bifid
The inner prong,
widely separated prongs.
bone,
further examina-
deeply grooved along
its
main body of the
whicli constitutes the
mesial border, producing two shelf-like projections
one above the other: these converge near the tip of the prong to form a slender pointed
style resting in a
groove on the outer border of the posterior extremity of the vomer as in
other Palwognathic, and as in the hemipterygoid of Neognatluv,
The outer limb
closely applied to the ventral
is
and external aspect of the extreme
hinder end of the maxillo-palatine process, extending as
which the maxillary portion of the quadrato-jugal bar
the outer and inner limbs the palatine
The palatine
seen
a
is
is
is
far forwards as the point
given
from above.
conceals trom above.
Its
like plate to articulate
has
its
from
In the cleft between
interposed, as will be seen presently.
It is closely applied,
throughout
backwardly projecting plate from the maxillo-palatine
this last
offi
short and broad, with a slightly bifid free extremity.
in its entirety
Thus
Impeunes. Colymbi.
e. g.
proximal extremity sends
off,
its
pi'ocess.
It
can only be
entire length, to
This
it
upward and inward, a
with, and underlie, the extreme
entirely
flange-
hinder end of the vomer.
right and left limbs sandwiched, on either side, between the
—
PHYLOGENY OF THE PAL.EOGNATH/E AND NEOGNATII^.
A
and the palatine below.
l)terygoid above
palatine helps to
fill
up the
cleft
mentioned above, the palatine
portion of this flange-like plate of the
between the pterygoid fork already described.
upon a backwardly directed
rests
palatine process, the whole of which
conceals from
it
As
plate from the maxillo-
the dorsal aspect.
Ventrally,
run along, and below, the external palatine border, and to
extreme posterior end. The forward extension of the palatine on
this process is
found
terminate near
its
to
the ventral surface corresponds with the level of the tip of the vomer.
is
205
This extension
derived from the ventral surface-layers of the main body, the palatine terminating
abruptly on the dorsal aspect in a V-shaped notch at the level of the middle of the
vomer.
This ventral process rests in a groove along the mesial border of the maxillo-
palatine process.
The
relations of the
say, that the free
vomer have already been more or
end of each posterior limb
is
less indicated.
wedged
Suffice
it
to
between the spine-like
in
extremity of the pterygoid above and the flange-like inward projection from the dorsal
surface of the palatine below.
The
palate of Apteryx will be found, on a close comparison with that of the remaining
Falwognathce, to have departed from the typical Struthious form in the direction of
that taken by Bltea, the relations between palatine and vomer being similar in both.
The ibrm
that
of the pterygoid differs from
of
all
other birds, in that
it is bifid, or,
in
sends off a separate limb whose mesial border passes downwards to underlie the
it
external border of the palatine.
The
sented.
itself,
tluxt
internal,
This external, palatine limb
elsewhere unrepre-
is
vomerine limb may be regarded as a flattened plate folded upon
The
the edge being inwards.
superior edge, or upper
representing the mesial border of an
must be regarded as
lip,
originally flattened pterygoid such as that of
Drovueus.
The dentary
Tinamous
it
of
is
great
The
which the ventral
spleidal
is
is
Caudad each ramus divides into
In Rhea the dorsal limb
the larger.
of great length in
all,
extending
jaw, and backwards to within a short distance of
quadrate.
Ii/iea,
Apteryx, and
extends backwards to within one-fourth of the whole length of the ramus.
It is relatively less in Struthio.
limbs, of
In Lrommis, Casuarius,
length.
It is
tlie
forvvaixls to
dorsal and ventral
is
paired.
the symphysis of
tlie
level of the glenoid cavity for the
rod-like in Struthio, laterally compressed in the remaining forms.
The atujular, in all save Apteryx, articulates, proximally, with the supra-angular,
being embedded caudad in its external border distally, it twists downwards so as to
;
form the ventral border of the ramus.
In Casuarius and Droma'us
and inner border comes into contact with the articular.
considerably short of
which
is
extreme posterior
Rhea and
Struthio,
hollowed out
proximally with
to receive
it.
the ventral border
The supra-angular
rests
dorsal border.
VOL. XV.
— PART
v.
it falls
this.
In Ajjteryx, the angular articulates
articular,
In
its
No. 13.
December, 1900.
2g
of
upon
the
its
ME. W.
206
The
sujn-a-angular, iu
border
P.
PYCKAFT ON THE MOKPHOLOGY AND
all,
proximally with the articular.
articulates
Its
dorsal
received between the right and left moieties of the superior limb of the
is
dentary.
ventral border descends caudad
Its
of the angular; distad
it
to
the
level of
the
border
inferior
rises gradually so as eventually to cross the dorsal
border of
the same.
The coronary
is
is
in all a short, slender,
many
In
and rod-like bone.
Neogiiatli(e it
often of considerable size and laterally compressed.
In
all
the Palceognatlm
it
appears
to
extend
backwards
so
as
embrace the
to
inner border of the articular and to form the internal boundary of the articular surface
In Ehea and Apteryx
for the quadrate.
described as vestigial.
very feebly developed distad, and
In Casuarius and Uromceus
In no case does
intermediate.
it is
it
it
is
much
may be
Struthw
larger.
is
extend forwards beyond the posterior third of the
ramus.
Such characters of the skull as are of use
found iu
ibr systematic purposes will be
the " Keys " at the end of this paper.
More
material
is
needed before these can be made complete, since a "
species has not been possible in
the case of Casuarius.
specific in the case of the other
considered to be
"
to the
the characters
genera will stand the
test of
time
In several instances I have been obliged to rely upon the characters
remains to be seen.
of a single skull, as in the cases of the skulls of
I feel justified in
Whether
Key
Ehea macrorhyncha,
for instance.
reviving Dr. Sclater's Struthio meridionaUs [90] on account of the
many marked peculiarities which the skull and the trunk of this skeleton presents.
The work of distinguishing the skulls of the four species of Apteryx has been a very
difficult
Indeed, as Mr. Beddard has already remarketl [8], " the very slightest
one.
differences, apart
from those of
Some Poikts of
The
size,
distinguish the Apteryges."
SIGNIF1CA^•CE ix the Struthious Palate.
following points appear to be worthy of special consideration and
may
fitly
be
discussed here.
In comparison with the palatines of the NeognathoB,
Rhea, as iu the other Paloeognathce, the
palatines
another by the laterally expanded feet of the vomer.
meet
iu tlie
it
will
be remarked that in
are widely separated
one from
In the Neognathoe the palatines
middle line, thrusting the vomer forwards, so that
articulate not with the pterygoid but with the palatines.
it
now appears
to
In Rhea and Crypturi we
way in which this has come about. Turning once agaiu
to the skull of Uromceus, we notice that the palatines are only loosely attached to the
outer border of the vomerine feet, whilst in Rhea (PI. XLII. fig. 5) the palatines have
have probably a clue
moved inwards
to the
so as to overlap these feet
and approach one another
mesially.
Their
a
PHYLOGENY OF THE PAL^OGNATH.^ AND NEOGNATH.^.
207
relation to the pterygoid has also changed, for they no longer are connected with
movement have come
outer border bnt by this same inward
In the Neognathv, the inward movement has attained
meeting one another mesially,
in so doing.
As
[82], this
not altogether
is
as
l)y
In
the case.
much
it.
maximum,
just remarked, thrusting the
a matter of fact, however, as
forwards in a spike-like form,
though but
we have
to underlie
its
the palatines
vomer forwards
we have already shown in earlier papers
the young bird the ])terygoid extends
the vomer,
as in lihcn, so as to articulate with
The
the slightest contact.
its
palatines have succeeded in
beneath these anterior pterygoid ends so as
to all
l)ut
moving inwards
the original
entirely sever
between them and the vomer.
relations
Later in
(soon after hatching) the severance
life
is
At
complete.
this stage, the
anterior end of the pterygoid fractures at a point corresponding witli the free end of
The
the palatine.
fracture later becomes a true joint, and the anterior end of the
pterygoid resting upon the palatine gradually merges with this bone so as to obliterate
of
all traces
materially from
the
that
free
pterygoid of
tlie
Neognatluv
is
a
the palato-pterj'goid connection in the late embryo not differing
that of the
vomer from the pterygoid
so
Thus the
original existence.
its
secondary feature,
palate of
Further, the apparent isolation of the
Fakcognatha'.
in the Neogiiathw is seen to be a cocnogenetic character,
group
this
brought into close relation with that of the
is
Palceognath(e.
A
further point of interest in this comparison between the Palaeo- and Neognathine
skull
is
the change which the
vomer
in the latter has
undergone in relation
])arasphenoidal rostrum, a change which indicates a shortening both of
to the
vomer and
rostrum.
The vomer
antorbital plate,
This
processes.
the NeogwiilKV rarely extends backwards beyond the base of
in
in the
is
Pahrognatlwi
may
reach nearly as far as
tlie
l>asipterygoid
an undoubted proof of the shortening of the vomer.
That the rostrum has
also
that in the Pahvognaflici'
the level
it
the
undergone a considerable shortening
it
is
shown by
tlie
fact
extends forwards for a very considerable distance beyond
of the lachrymo-nasal fossa, in
the Neognntha'
it
commonly ends
in
the
region of the antorbital plate.
Yet other evidences of
reveal themselves in
whilst these
and modification of the dromgeognathous palate
the Neognathw when we come to closely compare them, and
show how
shifting
closely the
two groups are related they show
still
more
tlie
lower grade of type persistent in the Pahrognatlue.
In the PaUfognatlim the free ends of the basipterygoid processes articulate with
the extreme proximal end of the pterygoid quite close to the articulation with the
quadrate.
In Neognathce these processes, when present, have shifted forwards on to
the rostrum, so as to articulate with the middle of the pterygoid.
The
forw^ard shifting of the
Neognathine vomer, which we have already noticed
2g2
—
208
W.
ME.,
which brings
shifting
plate
— appears
to
its
p.
PYCRAFT ON THE MOEPHOLOGT AND
proximal end more or
have been accompanied by a similar shifting of the palatines, since
main body of the pterygoid bone but with
these no longer are connected with the
we have
This, as
distal end.
by means of a joint
Neognathine
its
already shown, ultimately fractures and fuses with the
palatine, a joint forming at the line of fracture.
pterygoid
immediately under the antorbital
less
point of great difference between Palaeo- and
a
is
This connection of the palatine and
skulls.
In the Tinamidce we have an intermediate stage between the Patceo- and the Neognathw. The vomer is undoubtedly relatively shorter posteriorly than in Rhea, its
free
end lying midway between the
The
processes.
articulate with
and the basipterygoid
level of the antorbital plate
pterygoid has increased in length, so that the vomer and palatine
its distal
The quadrato-jugal
extremity only.
in length relatively, extending forwards
now beyond
fossa has also increased
the level of the antorbital plate
as far as the vestigial maxillo-nasal process.
A
study
careful
points
of these
will
greatly
facilitate
the conviction that the
segithognathous and schizognathous skulls are but modifications of the dromaeognathous
The desmognathous
type.
is
a further modification of the schizognathous palate.
The single-headed otic process of the quadrate in the Palwo(jnathw, upon which
so much stress has hitherto been laid, appears to have less importance than the points
to
which attention has just been draM'n,
for in A])teryx the otic process
is
two-headed,
as in Neognathce.
Thk Vertebral Column.
The memoirs
of
Owen, Mivart, and
struthious skeleton render
T. J. Parker on the vertebral
unnecessary to do
it
quite
a.
The Presynsacral
more than
column of the
briefly
comment
thereon here.
are heterocoelous,
presynsacral vertebras of the Palceognathoe
All the
this
Vertebrae.
region
are
free
except in
the
Tinamidce, in
and
all
of
which certain of the thoracic
vertebrae fuse.
The
vertebrae of iJromreus are the least specialized in type
;
those of Casuarius are
very similar.
Both
in
Dromoeus and Casuariits the cervical veitebrse are conspicuously shortened
In Casuarius the neural spine, traced from the head backwards,
antero-posteriorly.
undergoes considerable change of form.
Anteriorly
tubercle rising from the centre of a flat neural plate.
it
is
little
more then
a
median
At about the 6th vertebra
it
sends backwards a pair of Ioav ridges which terminate in a pair of hyperapophyses.
At about the 10th
crescentic form.
vertebra, the spine has exchanged its <-shape for a transversely
This, for the next 4-5 vertebrae, becomes broken across in the middle
PHYLOGENY OF THE PALJ<:OGNATH/E AND NEOGNATH^:.
209
form two high curved ridges sloping downwards and backwards to terminate in
metapophyses. At about the 15th vertebra the two ridges again coalesce and form a
line to
very high vertical transverse neural spine.
becomes more and more
backwards, bears a fossa at
The
vertebrae
very considerable
From 6-10
bone
to the
The
its
Every neural spine, from that of the axis
and
base, both anteriorly
The
posteriorly.
latter
from the middle of the neck backwards have these
They lodge
size.
the ISth backwards this gradually
laterally compressed, so as to pass insensibly into the typical
neural spines of the thoracic vertebrse.
deeper.
From
in C. casuarius
and
is
the
fossae of
a ligament.
C.
ausfralis the diapophysis sends back a bar of
c.
hyperapophysis.
vertebrae are all pneumatic.
In the hinder cervicals there
aperture dorsad of the interzygapophysial ridge.
very large ones
— one below the transverse
is
a large pneumatic
In the thoracic there are several
process, one
between the transverse process
and the postzygapopliysis, and one dorsad, lying between the anterior zygapophysis
and the base of the neural spine.
The
This
hist is
represented by a deep fossa in Lromceus.
aperture ventrad of the anterior zygapophysis
feebly developed or wanting in
is
Lromceus.
The
and liypapophyses resemble those of DronueuH.
cervical ribs (pleurosteites)
In Lromceus the vertebrae are
transversely
specialized
less
than
in
expanded neural spines are wanting, though
tlie
in front of the neural s[)iue.
by cancellated
The
is
very deep, as also
The
and a
longer than in Lromceus or
foss;e are
narrow grooves channelled out of
posterior cervicals have the neural spines wider and shorter, autero-
cervical ribs, as in Casuarius
As
much
and they are deeply hollowed for the ligament.
posteriorly,
these.
that lying at the base and
is
spines of the anterior cervicals are long, low, and rise to form
The ligamental
a sharp median ridge.
The
fossa in, or
tissue as in Casuarius.
The neural
this ridge.
fossae,
sides of the fossae, moreover, are smooth, not obscured
In Struthio the centra of the vertebrae are relatively
Casnariits.
ligamental neural
The pneumatic
especially that caudad of the neural spine, are very deep.
above, the interzygapophysial ridge
The high
Casuarius.
in
Lromceus
tliey fuse
lateral parapophysial
the prezygapophysis.
vertebral artery.
One
and Lromceus, are long, but more slender than
in
with a plate of bone depending from the diapophysis
outgrowth from the anterior end of the centrum below
The presence of
this rib
serves
great point of difference between
to
tliis
enclose a canal for the
region of the vertebra and
that in Lromceus and Casuarius lies in the fact that in ISfruthio the lamella depending
from the diapophysis, and with which the
the centrum
for a considerable distance,
rib articulates, is
continued backwards along
forming an extensive and tunnel-like passage
for the artery.
The pneumatic
apertures are not so conspicuous as in Lromceus.
There
is
aperture in the interzygapophysial ridge of the anterior cervicals as in Lromceus.
no
In
ME. W.
210
the hinder
and
cervical
P.
PTCRAFT ON THE MOEPHOLOGT AND
thoracic
vertebrse
there
is
a
large aperture beneath the
in the roof of the tunnel for the vertebral artery, or in the thoracic
diapophysis, opening
vertebra below the prezygapophysis and between the dia- and parapophysial articulations of the tuberculum
The deep
and capitulum.
fossa at the base of the anterior
border of the neural spine of the hinder thoracic vertebrae
In Mhea the vertebrae have departed
are very long and cylindrical.
The
vertebrfe.
still
here very shallow.
is
more from the normal
The
type.
cervicals
Neural spines are present only on the 2nd
to 5th
free hinder borders of the postzygapophyses of the 6th vertebra are
received into a pair of shallow pits on the 7th, lying mesio-caudad of the prezygaCervical ribs are only feebly developed.
pophyses.
From
the 2nd to 11th vertebrae
they are represented only by short styliform processes; from this vertebra backwards
they are represented only by the extreme anterior articular end of the rib which serves
to
As
form the band-shaped external wall of the vertebrarterial canal.
canal
vertebrarterial
is
roofed by a backward extension
of the
in Struthio, the
lamelliform plate
depending from the diapophysis and serving as an articulation for the cervical rib.
This plate is described and figured as the " rudimentary rib " by Mivart [63]
doubtless by a
slip.
In Ajiteryoc the form
The
Bronunis.
rest
of
of the atlas and axis vertebrae bears some resemblance to
its
The neural
herewith associated.
On
spines never bifurcate.
they form compressed blades from the 8th to the 12th
vertical in
from those of the forms
vertebrae differ conspicuously
;
the anterior vertebrae
they are columnar and
form and central in position with regard to the neural laminae.
diapophyses of the 6th and 7th vertebrae project outwards and
the level of the prezygapophysis;
wards
into
;
from the 7th
to
The
backwards beyond
the 13th backwards and down-
from this vertebra caudad they project directly outwards, and merge gradually
The neural
processes of the thoracic vertebrae.
the transverse
spines of the
thoracic vertebrae are equal in size, of great breadth antero-posteriorly, and interlock
along the dorsal ridge by anterior and posterior bifurcations as in
There are no pneumatic apertures
many
Neognatlice.
Cervical
in either cervical or thoracic vertebrae.
ribs are vestigial.
In Crypturi the thoracic vertebrae are anchylosed and bear hypapophyses.
these occur only
Palceognathxe
penultimate
ridge
is
is
free,
on
the
posterior cervicals
the ultimate fused with the synsacrum.
perforated by pneumatic apertures.
In
the
(cervico-thoracic).
In the
The
The interzygapophysial
cervical
vertebrae
pneumatic
apertures open into the roof of the vertebrarterial canal.
The diapophyses of the cervicals project forwards beyond the prezygapophyses.
The neural arch is constricted in the middle, immediately behind the prezygapophyses.
The vertebrae of Dinornis resemble those of Casuarius. The fossae lying beneath
the transverse process of the thoracic vertebrae, absent in Bromceus, were much larger
relatively
than in
Casuarius.
The
cervicals
resembled
those
of some
species of
PHTLOGENY OF THE PAL^EOGNATH^ AND NEOGNATH.E.
Casuarius in having a bony bar running
from
211
diapophysis backwards
tlie
the
to
hyperapophysis.
The
thoracic vertebrm call for no special remark here.
They have already been
minutely and carefully described by Mivart.
The
known
vertebrae
backward
probably be regarded as true
shifting of the sternum.
Writing
T. J. Parker.
late
as the cervico-thoracic should
the ribs of which have lost their sternal segments through the
thoracic vertebra?,
This view has already been promulgated by the
" The occasional absence of
of Apfcrz/x, he says
:
the cervico-thoracic uncinates, taken in connection with the vestigial condition of the
one case and with the fact that in the same skeleton the
ribs in
first
thoracic rib
of the right side terminates ventrad in a blunt free end and has no sternal portion,
seems
to point to
atrophy of their
an inclusion of anterior thoracic vertebrte
the cervical region by
in
ribs."
b.
The Synsacral Verfebm.
In Casuarius there are 8 (eight)
[)re-
(PI.
XLIV.
fig. 5.)
and 8 (eight) post-sacral
vertebrae,
with
tlie
two true sacrals numbering eighteen synsacrals.
Of
2nd
these,
to
two are thoracic.
The 1st lumbar has large parapopliysial processes the
much smaller; the 4th and 5th become larger and
;
ord of this series are
abut, the 4th against the pectineal process and the 5th against the ischium.
these follow two lumbo-sacral.
The
Behind
true sacrals are closely crowded and have slender
parapopliysial processes, which abut against the ilium and ischium immediately behind
Behind these in
the acetabulum.
licarunculains
C.
parapophysial processes abutting against the ilium.
might readily be mistaken
which follow
processes.
six
They
for true sacrals.
two pseudo-sacrals with large
lie
But
for the sacral plexus
are really the
iirst
they
two caudals, behind
fused caudals with diminishing and backwardly directed transverse
In other Cassowaries there
is
The
but one pseudo-sacral.
sacral vertebrae
do not bear diapophyses.
In
Dronmus
the
number
of synsacral vertebne
is
the
same
The
as in Casuarius.
6th lumbar, however, does not send out a parapophysis to abut against the pectineal
process.
The neural
spines of the synsacral vertebrae are all highly pneumatic, and
They remain separate
of loose cancellated tissue.
in the adult are fused into
young bird
the
one mass,
pre-sacral
all traces
vertebrae
are
separated, so
that
to
some time
after hatching,
of separate vertebrae being
just
distinguishable
approximated pre-acetabular legious of the ilium.
spines laterally expanded so as
for
The
made
lost.
u[)
but
In the
between the closely
post-sacral
have the neural
keep the post-acetabuhir ilium moderately widely
is
a deep loculus (PL XLIV. fig. 5).
between each vertebra
In the adult the whole becomes roofed over by a thin bony
from the post-acetabular ilium of one side
to that of the other.
plate extending across
MR. W.
?J2
P.
PTCEAFT ON THE MOEPHOLOGY AND
even three-quarter grown birds the neural spines from
in the half or
Seen dorsally
above the acetabulum backwards
for
some considerable distance are found
take
to
the form of vertical transverse plates, forming a series of loculi between the post-ilia.
Thouoh
further back in the series the transverse plates
In the adult
remain.
become columnar, the
loculi
loculi are completely covered in by a thin sheet of bone,
tliese
formed by the fusion of a series of thin plates capping the neural spines.
In
the
embryo Bromoeiis
XLIV.
(PI.
fig.
6)
it
worth noting that the
is
lofty
transversely expanded neural spines of the sacral and anterior post-sacral vertebrae of
the adult are i-epresented only by low cartilaginous ridges, and bounded on either side
In the dried skeleton the carti-
by the free edge of the dorsal border of the ilium.
laginous portion shrinks up, revealing (1) a pair of ossified neuroids, which
meet one another
middle
the
in
line,
and
The
the similar cartilaginous ridges already described.
foundation of
height of the neural spines
to
low osseous ridge marking the
a
(2)
fail
increase in
followed by a corresponding increase in depth of the
is
ilium (p. 227).
The
centra of these synsacral vertebrae in this early stage are, in the dried skeleton,
represented only by a
is
The
stirrup-sliaped.
much
bony
flat
plate, so that a single vertebra, with
This
neural arches,
makes the neural space of a relatively
The nerve-apertures are single and not paired
thinness of the centrum
greater size than in the adult.
as in tlie adult.
its
is
shown
well
in PI.
XLIV.
figs.
5-6.
In Struthio there are ten pre-sacral and eight post-sacral vertebrae.
The
series
lumbar
commences with the
two thoracic
The parapophysial elements
vertebrae.
The diapophyses
last
of
tlie
vertebrae.
of these
are
on the 4th, they gradually increase in length up to
On
rod-shaped bars of bone.
vertebrae
vestigial parapophyses.
The
left side
the 10th, this process
ai'e
short and
The
last
Museum
lumbar bore
extended backwards to join
sacral vertebrae are
two
in
where they form long
short and rod-shaped.
8th,
tlie
is
Short and triangular
collection the lumbo-sacrals
distinct parapophysial elements,
tlie first sacral rib.
number and have
large diapophyses.
They
caudad than those of Casuariun, resting against the hinder border of the
ischiadic or post-acetabular process of the ilium,
of
the
acetabulum.
acetabulum.
The
thick.
possibly represented by a single vertebra which bears
In one skeleton in the
were certainly unrepresented.
which on the
all
follow seven
lumbar vertebrae from the 4th backwards extend upwards
on to the inner side of the ilium as forwardly directed spikes.
The lumbo-sacral
Then
In
Casuarius
the
sacral
lie
more
downward
and therefore conspicuously caudad
ribs
lie
immediately behind
the
outer free ends of the sacral ribs give rise to a large swollen process
from the dorsal surface, which extends upwards and backwards to reach the diapophysis
of the 2nd sacral, sometimes extending forward to that of the
in the
the
Museum
left side.
1st.
In one specimen
collection the lumbo-sacral has developed a parapophysial process
on
—
:
PHYLOG^ENY OF THE PAL^OGNATH.E AND NEOGNATH.E,
The
1st caudal bears
those of the sacral.
free caudal
a'^
is
both para- and diapophyses.
Behind
encountered.
this lie seven caudals
The neural
213
Sometimes the former fuse with
without parapophyses, before the
first
spines of these are not transversely expanded
in Casuariiis.
In Rhen there are ten pre- and
The
pre-sacral comprise two thoracic, four lumbar,
of these
The
five post-sacral vertebrae.
may develop parapophysial
saci'al
and four lumbo-sacral
the last
processes.
ilium more dorsad
ribs fuse with the
;
than in Struthio, Casuarius, or
In the last three they abut more or less extensively against the post-acetabular
Bromcevs.
ascending process of the ischium.
In Wiea they
rise dorsad, so as to
reach the level
of the centre of the acetabular arc.
The
1st sacral is intermediate in position, with regard
Casuarins and StnitJiio.
In the
The
sacral
ribs
are
very
be remembered,
latter, it will
the acetabulum, in Casvarius level with
short
its
to the acetabulum,
it lies
considerably behind
hinder border.
and almost indistinguishably fused with their
In the possession of diapophyses these vertebrte
diapophyses.
between
differ
from those of
Casuarius and Bromceus, and resemble Strutldo, in which, however, they are
much
more extensively developed.
The
first
The 2ud caudal bears diapophyses
but they are almost indistinguishably fused.
The
para- and diapophysial processes,
post-sacral, or 1st caudal, bears both
only.
3rd to 5th caudals are somewhat remarkable in that they are represented only by
atrophied vertebrae.
The centra
of these are reduced to thin delicate rods, supporting
slender diapophyses and feeble neural spines.
The 6th vertebra resembles the
but has a stronger neural spine.
vertebra to escape from the clasp of
It
the
is
The 7th has
the post-acetabular ilium.
a
first
much
thicker centrum, but like the 6th has lost
inflated cancellous centra
with the ischia.
its
stronger neural spine and slightly
diapophyses.
and short neural spines
The 10th
is
the
first
5th,
;
The 8th and 9th have
they fuse by their ventral aspects
which there are eight
free caudal, of
in all
these have short thick centra and short neural spines.
In Apteri/x there are nine pre- aud
sixteen in
The
first
five
or six post-sacrals, with
the true sacrals
all.
pre-sacrals represent
two thoracic, four lumbar, and three lumbo-sacral.
post-sacral (1st caudal) bears short para-
and diapophysial processes.
The
The
post-sacral
are chiefly remarkable for the great lateral compression which they have undergone.
The synsacrum
of the Dinornithido' differs in
other Palceognatlice.
It
resembles in
many
many ways
respects
from that of
that of DromcBus
all
the
and Casuarins
on the one hand, and that of Struthio on the other. The pre-sacrals are apparently
There is an
eleven in number, viz., two thoracic, six lumbar, and three lumbo-sacral.
indication of a fourth lumbo-sacral bearing parapophysial processes, which have almost
completely fused with those of the 1st sacral.
VOL. XV.
—PAET
V.
No. 14.
December, 1900.
There are
five post-sacrals.
2 h
ME.
214
P.
A^".
PYCBAFT ON THE MOEPHOLOGT AND
pre-sacrals resemble those of
The
in that they descend
Brommis and Casuarius
below the level of the ventral border of the pre-ilium, but not greatly so. lu Strufhio
this feature is much more marked, the whole centrum and base of the neural spine
being exposed
in
side view.
directed backwards instead of
The parapophysial processes
From this cause
forwards.
of
the
caudal are
first
the true sacral
vertebrae
are completely isolated.
In Stnithio, Casnarivs, Dromcens, and Apterijx, it will be remembered, the para
pophysial process of the first or even first and second caudals may combine with
those of the true sacrals to form a sacral mass, the true nature of which is difficult
The post-sacrals (caudals) resemble those of Strutkio
to make out in the adult.
rather than any other Fafito' in the great development of the diapophysial process,
which extends upwards, outwards, and backwards to separate the post-acetabular
ilium.
In Binornis, however,
this
separation of the post-acetabular
The neural
greater length than in any other Ratite.
ilia
carried
is
to a
spines of the BinornitUda' are
peculiar in that they combine dorsally to form a broad lozenge-shaped superficial plate
of bone which serves as a cover to the
huge
fossa that otherwise exists
upwardly directed transverse processes and the post-ilia.
The synsacrum of the Crypturi differs from that of
Moreover, in the points wherein
described.
it
differs
all
between the
the other forms
from these
it
herein
resembles the
Neognathce.
One
neural
of the most noticeable diff"erences
spines.
In
all
is
the fact that the post-sacrals have lost their
the flightless Pakeognathae these are of great length.
diapophyses of the post-sacral region
rise
The
from the top of the centrum, at the base of
a long, faint, median ridge marking the position of the neural spines in other forms.
These diapophyses develop thin plates of bone which fuse one with another to form a
lozenge-shaped plate, protecting the inter-diapophysial
fossse, as is
done by the similar
plate from the neural spines in Binornis.
The synsacrum
and
five
pectineal
of the Tinamous includes one thoracic, four lumbar, five lumbo-sacral,
post-sacral.
process
as
There are no parapophysial processes abutting against the
This process, it should be
in the flightless Pakeognntlm.
mentioned, has shifted further backward relatively to the sacral vertebrae, so that a
line drawn across from the base of the anterior concave border of the one would pass
immediately in front of the
The
first sacral.
sacral ribs are very long, and articulate with the ilium immediately behind the
acetabulum by a long
vertical
roughened
surface.
The
rib
of the second sacral
is
Aestigial or wanting.
The
free caudal vertebrae in tlie Palceognathce
times the 9th and 10th are more or
In Struthio the 10th
is
laterally
do not exceed ten in number.
less fused.
compressed
to
form a kind of pygostyle.
Some-
PHYLOGENY OF THE PAL.EOGNATH.E AND NEOGNATH^.
215
In Bhea there are but seven free caudals. This
of the ischia.
Counting from the
is due to the great backward extension
end of the post-acetabular ilium backwards, as
free
in other forms, tliere are eleven post-iliac caudals.
Ribs.
The
It
to
anterior cervical ribs (pleurosteites) have already been described.
probable that the free cervical ribs (cervico-dorsal, Mivart) and the vertebrte
really belong to the thoracic series.
This is indicated by the
is
which they articulate
that these ribs have lost their connection with the sternum, and have lost their
fact
sternal ribs through the
In Casuariiis and
and Rhea, two.
a shorter and
backward shifting of the
Lromaus
there
may be
sternal plate.
three or four free cervicals.
In Struthio
In Apteryx one, which bears a large uncinate
more
style-like rib anterior to
tliis
(in A. oweni there
and wanting the uncinate).
is
In the Crypturi there are two free cervicals.
The
Bromaus
thoracic ribs in Casuarius and
agree with Struthio, and differ from
Rhea, in the fewness and slight development of the uncinates.
In Casuarius the 1st
of sternal
The 6th and
ribs.
more or
pair are
five pairs
less
7th are long, but have no sternal segments.
overlapped by the pre-ilium.
underlie the pre-ilium.
The 9th
In
little
C. hennetti
long sternal segments, but they
means of
fail to
however, that they have not long
The uncinates
appear
to
grown C
pair are very short
and
scale-like vestige reprein the
young bird can
pre-ilium.
reach the sternum.
lost their
may
most of the skeletons which
I
In a
These have been probably
lost in
half-
maceration
have examined.
articulates with the anterior lateral
;
the second
is
process of the sternum.
very
long,
and almost
Following these come
four pairs of thoracic ribs connected with the sternum by sternal ribs.
succeeded by a pair (7th) with very long sternal segments, but which
The next two
this,
Generally only two or three
anchylose with their respective ribs.
salvadorii I find five uncinates.
very
connection therewith.
of the ribs of Casuarius are never large.
be present, and these
The oth pair have
We may gather from
sternal segments.
In JJromu'us there are two free cervicals
sternum.
The 7th
there are but three free cervical ribs and only four pair of thoracic ribs
articulating with the sternum by
in
The 8th
pair are represented by a
more than the tuberculum and capitulum, which
be made out {e.g. 1374 a Mus. Coll.) closely applied to the
senting
still
of thoracic ribs articulate with the sternum by means
pairs have no sternal segments,
fail to
These are
reach the
and are overlapped by the
prr-ilium.
In Struthio there are two free cervicals and five pairs of thoracic ribs articulating
with the sternum by means of sternal
sternal
ribs.
The
pair next behind these has long
segments which do not reach the sternum.
collection of S. camehis this Gth sternal
In a mounted specimen in the
segment actually articulates with the sternum.
2h2
ME.
216
PTCEAFT ON THE MOEPHOLOGT AND
pair (7th thoracic) are long, but without sternal segments
The following
pair are short
"W. P.
;
the 8th
and fuse with the pre-ilium.
In Rhea there are two free cervicals, three pairs of thoracic articulating with the
sternum by sternal ribs, and provided with very long uncinates. The 4th and 5th
pairs are long, but have
The 6th
no sternal segments.
pair are very short and fuse
with the pre-ilium.
Amongst
some
Dinornithidce there appears to be
In Dinornis maximus there are three free
variation.
cervicals,
sternum by sternal segments, one long free
)ib,
two thoracic connected with the
and three
free ribs beliind this
overlapped by the pre-ilium.
In Pachyornis there seems to have been but one free cervical or cervico-thoracic,^
three thoracic articulating with the sternum, four long free ribs, and short free ribs,
The uncinates were very
these last two being overlapped by the ilium.
long.
In Apteryx the ribs are remarkable for the great breadth and flatness of their
There may be one or two pairs of cervico-thoracic
vertebral segments.
there are four
;
thoracic articulating with the sternum (in a skeleton of A. australis I found five on
one
side,
and
in
a skeleton of A. oweni, both in the Rothschild Coll., I found but
three thoracic vertebrae connected with the sternum) and four pairs of free ribs behind
these,
all
of which
The
by the pre-ilium.
overlapped
are
Six pairs of ribs bear uncinates, the series
exceedingly short.
The uncinates
last cervico-thoracic.
are
often
commencing
in the
last
pair
are long and broad, extending backwards to the
hinder border of the rib next behind.
In the ribs of
the Fakeognathce, save Apteryx and some Tinamous, there
all
is
a
large pneumatic aperture lying in the cleft between the tuberculum and capitulum.
The Sternum and Pectoeal Arch.
a.
As with
The Sternum.
the other portions of the skeleton so far examined, so with the sternum,
JJronueus possesses the most generalized type.
The sternum
of
the young
similar in appearance
;
Brommus and
that of the
young Casuarius
are very
in the adult they are readily distinguishable.
In the adult Dronicens the anterior lateral processes are very long, slender, and
directed upwards and backwards.
emargination in
developed.
length
The
their
place.
articular
The
differ
no spina externa nor interna, but a deep
is
ventral
surfaces for
the distance between the
These grooves
There
lips
the
articulation
from those of
all
of the
coracoids
for
the
coracoid
are
first
short,
grooves
not
and third
are
sternal
in
ribs.
the other Palceognathce in that they overlap,
causing the bases of the coracoids to cross one another in the middle line as in
Carinatce.
well
exceednig
many
PHYLOGENY OF THE PAL^OGNATH.E AND NEOGNATH^.
Fig. 4.
B,
A.
\
/
V-
a^//!
r
1
^t
f
/
—W^/'-''-
'^^-lls^-
c.
i
^-y
.-^'
\
«!•.
<^
•^^
217
ME. W. P PYCRAFT ON THE MOEPHOLOGT AND
218
The sternum
common
As
in
of Dromceus ater
is
remarkable, possessing as
with both Casuarivs and Rhea, but
I),
at the
is
does cliaracters in
it
same time Dromseine.
noire-hollandice, the anterior lateral processes are conspicuously largo
have the same form.
The
outline of the free edge of the corpus sterni from the
articulation of the last thoracic rib
That of the right
are obsolete.
and
the lower lip of the groove.
side
backwards
is
This, as
The
the same.
is
represented by a portion
is
(jf
coracoid grooves
the inner angle of
seen in the figure (4 b), persists in the form
The obsolescence of these grooves indicates that
the pectoral arch and wing were much more degenerate than in Dromceus novrt^The corpus sterni differs from that of D. novw-liollandke in
hollandiw or I), irroratus.
the relatively much greater development of the protuberantia sterni, which resembles
of a median asymmetrical tubercle.
that of Bhea, and in the relatively narrower width across the region of the incipient
posterior
lateral
Four
processes.
ribs
articulate with
sternum,
the
in
D. novce-
hoUandia' only three.
The following measurements
in millimetres are of interest
B.
novce-Jiollandice.
Greatest lengtli
Greatest width (across aut.
Least width (across post.
It
should be
indicating a
.
J),
ater.
95
10
90
105
55
1
.
D.
160
...
lat. proc.)
noted that the metasternum of
ater bears
traces
its
length.
In Casuarius
articular surfaces for the sternal ribs are divided
it
is
much
of fracture,
The breadth
somewhat greater length than the above measurements.
the sternum nearly equals
The
lat. proc.)
:
of
longer than broad.
one from another by pneumatic
apertures forming pockets of great depth, extending from Avithout inwards towards
the middle line.
No
The hinder
border, both in Dromcpus and Casuarius,
external lateral processes are present.
The
sternal plate
is
is
shield-shaped.
of great
thickness and
highly pneumatic.
In the young Dromceus the anterior lateral processes are very short, and the ventral
margins of the coracoid grooves are almost confluent and not separated by a deep
median notch.
In Casuarius the sternum
is
relatively
much
longer than in Dromceus, the increase in
length being most noticeable from the articulation for the last thoracic rib backwards.
The
coracoid
obsolete.
grooves are
much
longer, and
the anterior lateral process
is
almost
In the middle line between the ventral lips of the coracoid groove in
Casuarius there opens forward a large pneumatic aperture
;
this
runs backwards into
the body of the bone for some considerable distance, penetrating a low median ridge
or protuberance resembling a vestigial keel.
a shallow groove taking
its
place, the sides of
raised to form a pair of long low ridges.
This
is
wanting in C. unappendiculatuSf
which further back become conspicuously
PHTLOGENT OF THE PAL^OGNATH^
The
219
sternal plate of Casucaius
Such
is of great thickness and highly pneumatic.
have been able to find will appear in the " Key."
resembles that of Lronums and Casuarms in several respects,
specific differences as I
The sternum
but
A]ND NEOGNATH.E.
of
Mea
yet very readily distinguishable from either.
is
The coracoid groove, though
shallow,
is sharply defined.
The outer and inner lips,
though but feebly developed, are subequal in length. The groove of the left side is
separated from its fellow of the right by a smooth-lipped and gently hollowed plate
of bone representing the anterior border of the sternum, equal in length to the groove
itself.
The anterior lateral processes are large.
The
articular surfaces for the ribs
are
behind' the anterior lateral process.
Thus
the distance from behind the articular
surface for the last rib to the hinder border
space separating the coracoid grooves
ribs
occupy only one-sixth of the whole
more than twice the distance of the
is
other words, the articular surfaces of the
or, in
;
and follow immediately
closely crowded,
lateral sternal border, in
Casuarius
it
occupies
about one-third.
The body
of the sternum
notched posteriorly and tapers from before backwards.
It presents a deep basin-like hollow dorsad and is i)roduced ventrad into a Ions,
is
The corpus
swollen, ridge-like prominence.
is
much
thinner than in Bromceus
articular surfaces for the sternal ribs are each
more
or less distinctly divided
sterni
or Casuarius.
The
into two, the outer being the larger.
The
pocket-like interarticular spaces (incisurse
intercostales) are of great depth.
Large pneumatic apertures open below the base of the anterior
further remarks on the sternum of Ehea, see p. 224.
In Struthio the sternum
Uromaus
in
is
roughly about as broad as long.
some respects and Ehea
grooves resemble that o{ Brommis.
Both dorsal and ventral
lips
in others.
The
The form and
ventral lip, however,
converge mesiad.
The
lateral process.
It
resembles
position of
is
its
For
that of
coracoid
wider than the dorsal.
right and left grooves are separated
by a slight hollowing out of the anterior sternal border.
The
anterior lateral processes
are moderately well developed and project outward and forward.
The
width and occupy nearly the
border of the sternum, thus differing from both Bromceus and Bhea.
pocket-like pneumatic incisurse intercostales are relatively less deep than in the
whole
The
articular surfaces for the sternal ribs are of great
lateral
foregoing forms.
The median
portion of the bar serving for the articulation of the
ribs has lost its glenoid surface, wjiich
Struthio and
Bhea
The
is
preserved only at either end.
In this point
agree.
There are large posterior
and
is
lateral
ventral prominence of
tlie
and
in
some sterna incipient median
sternum
lies
broader and more flattened than in Rhea
middle of the sternal plate.
lateral processes.
behind the centre of the sternal plate,
;
in
which, by the way,
it
lies in the
ME. W.
220
PTCEAFT ON THE MORPHOLOGY AND
P.
Apteryx, in the form and position of
coracoid grooves, resembles Rhea more
Like Rhea they are widely separated by a deeply
any other group.
closely than
its
concave median portion of the anterior border of the sternal plate.
the base of the anterior lateral process.
It
is
The groove
lies at
very short, and has a conspicuously
projecting lower lip.
The
from that of
sternal plate differs
breadth
Its
exceeds
all
the foregoing in being non-pneumatic.
Posterior lateral processes are well marked, and
length.
its
The
divided one from another and a median process by a pair of notches.
articular
Struthio, extend along the greater part of the lateral
surfaces for the ribs, as in
sternal wall.
Occasionally,
Beddard
e. g.,
in an
A.
descrilies a pair in
austrab's described
by Owen, the sternum was fensterated.
In another of
A. australis mantelU.
species were a
this
pair " of pin-hole size."
Parker directs attention
As Beddard remarks,
A.
haastii,
more
and
I
to a distinct keel
tliough not easy to see, this
think to these
easily seen, often occurs
The
on the sterna of A. australis mantelli.
is
we might add A.
readily
He
felt.
A
found traces in
similar keel,
and much
able to confirm,
show that
oweni.
on the sterna of Struthio.
researches of T. J. Parker and Beddard, which
I
am
can be obtained from the sternum of Apteryx.
Roughly, as Parker and Beddard point out, " in A. australis the length of the
no really reliable
specific characters
corpus sterni appears to be constantly more than half its breadth."
In A. australis mantelli " the length of the corpus sterni is often considerably
—
less
than half
its
breadth."
In A, oweni " the length of the body
is less
than half
its
breadth."
A. haastii, according to Beddard, belongs to the same category as the
The emargination
metasternal and posterior lateral processes vary so
upon as
The
last two.
of the anterior border of the sternal plate and the length of the
much
that they cannot
be relied
specific characters.
anterior emargination of the sternum seems to offer at least one
Thus, both Parker and Beddard agree that this
constant character.
but that A.
semicii'cular incision,
atistralis
mantelli
may be
more
is
or less
usually a
distinguished " from
A. australis by the greater depth of the emargination in the former subspecies."
Mr. Beddard
finds at
least
one exception
points out that " in A. oweni
....
to
this,
in
A.
sinuous, each side presenting a sigmoid curvature."
Parker
a^istralis mantelli.
the emargination of the anterior border
There would appear
slightly
is
to
be no
exceptions to this rule.
The sternum of
Dinornithidce more closely resembles that of Apteryx than of any
other form herein described.
From
this
it
differs
in the
following points
:
—That
portion of the anterior border of the sternal plate which separates the coracoid grooves
in
Apteryx
is
in the present group perfectly straight instead of hollow.
Furthermore,
—
PHTLOGENT OF THE PAL.EoaXATlLE AND
much
this region of the sternal plate is
The coracoid grooves
upwards.
Apteryx, these grooves
The
tliickcned to form a prominent lip, looking
at the antcro-lateral angle of the
lie far apart,
processes are of great lengtli
The form
different
groups into which
sternum
tlie
is
in
sternal plate.
lie
The
process, as in Rhea.
metasternam
Dinornithes are divided.
Apteryx does not extend beyond
in
;
lateral
of this and of the posterior lateral
tlie
As
the articular surfaces for the sternal ribs
;
crowded together immediately behind the anterior
posterior lateral
221
are exceedingly sliallow and only just visible.
anterior hxteral processes are short
length.
XEOtiXATII.E.
often of great
much
processes varies
The length
in the
of the meta-
posterior lateral processes.
tlie
In ^pyoniis the sternum resembles that of Apteryx more nearly than any other
Pakeocjnatha'
As
.
Apteryx,
in
the anterior border of the sternal plate
is
notched, the coracoid grooves are widely separated and very similar in form.
facets are
wide and well spaced along the
together as in lihea or Dinornis.
processes, but there
is
deefily
The
lateral border as in Ajjferyx, not
crowded
There are short but well-marked posterior
lateral
As Andrews has pointed out
no metasternal element.
rib-
in a
recent and admirable paper on the skeleton of this bird, the sternum of uiEpyorniti
represents an embryonic stage.
As Mr. Andrews has pointed out
[4],
the " sternum of JEjyyorms consists of the two
primitive costo-sternal elements only, and in this respect corresponds to an embryonic
stage in the develo])ment of the sternum in the recent Ratitte."
this
seem
to apply,
The sternum
he goes on
to point out, to the
the Crypturi differs from
of
chiefly in the possession of a large keel.
of Anomalopteryx
in
This
.
is
chiefly
due
spaces
crowded and
—
on either
incisurse intercostales
the sternal ribs
is
embryonic sternum of Apteryx.
that
of
the
flightless
Pakeoqnatlice
to the large posterior lateral processes,
which
articular surfaces for the sternal ribs are
side of the coracoid grooves.
—form deep pocket-like apertures,
The crowding together and forward
for instance.
The
lie
especially does
In general form the corpus sterni recalls that
The
Cryptnri arise very far forwards.
closely
More
The
interarticular
as in Dinornis
and Ehea,
position of the articular surfaces for
another Rheiue character.
anterior lateral processes are very similar in form and position to those of Rhea.
which they
still
further resemble in that the ventral aspect of the base lodges
a
pneumatic aperture.
The
coracoid grooves in the
the corpus sterni.
lips are better
IS
Crypturi are se[)arated by a deep gorge carved out of
These grooves
differ
from those of Rhea in that the upper and lower
developed, and in that their transverse extension
is less
while their depth
greater relatively.
There
is
a large spina interna.
Turning once again
kindly lent
siguiflcance.
VOL. XV.
to the
corpus sterni and comparing that of Rhea macrorhynchu
me by Mr. Beddard,
To
— PART
with that of a
wit, that in Rliea
v.
No.
15.
on
Tmamou. we
eithe]- side
December, 190U.
notice a point of
some
of the great mci\.mii protuberant iu
2
i
ME, W.
222
PTCEAFT ON THE MOEPHOLOGT AND
P.
near the middle of
sterni,
account of
Now
thinness.
its
extent
its
4
(fig.
c),
is
an area of bone conspicuous on
seems more than probable that these thin areas are
it
the last traces of a once deep fissure which has gradually closed up, thus uniting to
Parker has noticed a
the body of the sternum a pair of posterior lateral processes.
similar indication of the closui'e of these fissures in Apteryx.
Fusion probably
took place, in the case of Rhea, between the free ends of the
first
processes and the mesosternum by syndesmosis of the
A
invariably closed.
such as
is
is
fissure is
and raetasternum,
this process of closing.
It
generally thicker posteriorly
would contribute towards the work of obliteration. If, once
we imagine these fissures open, the resemblance between the sternum oi Rhea
and Cryptnri
It has
;
this also
be found to be significantly striking.
will
Gadow
been alieady suggested by
a degenerate keel.
is
would materially hasten
be noted that the membrane closing this fissure
also to
than anteriorly
again,
slight expansion of the free ends of the processes
indicated in Tiiiamidoe,
is
membrane, by which the
that the protuberantia sterni
This seems quite probable.
Seen in
quite as close as that claimed for Aptertjx by Parker
so conspicuous in the living bird
comus and
is
4
represent
resemblance
The sternal
Neognathw in
callosity
d).
Opistho-
a quite secondary character.
The Pectoral Arch.
b.
The
(fig.
represented amongst the
is
may
profile its keel-like
shoulder-girdle
of
the
Pala'ognathce
flightless
has undergone
The coracoid and scapula
modification and bears evidence throughout of retrogression.
are no longer free, except in the very
young
bird,
considerable
but completely anchylosed, forming,
the one with the other, a very obtuse angle, instead of an acute angle as in
modern
birds.
In Droriutus the sternal (epi-coracoid) border of the coracoid, though wide,
the length of the whole coracoid.
the coracoid grooves
The pro-coracoid
several
The scapula
is
ligamentous.
slender,
and looks
It bears a strongly
is
The
free
this border
is
than
received within
and underlies the anterior-lateral process.
supra-coracoid foramen
is
large and receives
like
an outward, upward, and backward outgrowth
marked
articular f;xcet for the vestigial furcula.
pectoral arch of Dromceus ater does not appear to differ materially from that of
D. novm-hollandioe.
made
;
Only the inner half of
outer portion
less
pneumatic apertures.
of the coracoid.
The
is
its
is
for
me
at
I
gather this from a sketch by Dr. Henri P. Gervais, kindly
the request of Prof. Oustalet, to
whom
I
am much
indebted, as
well as to Dr. Gervais.
The following measurements
(in millim.)
Length from pro-coracoid process
accompanied the sketch of Dr. Gervais
to tip of scapula
0"127
Greatest length of coracoid
0-062
Width, epi-coracoid region
0-047
:
PHTLOGENT OF THE rAL.EOGNATH.E AM) NEOGNATILE.
In the sketch
the
supia-coracoid foramen
not indicated, neitlier
is
223
is
there any
indication of clavicles.
In Casuaritts the coracoid
equals
its lengtli.
Dromceus,
tlie
relatively
is
much
whole sternal border
I'lie
pro-coracoid
is
is
form and
In Ehea the coracoid
Unlike
In the
a small pro-coracoid fenestra.
is
The
ligamentous.
snpra-coracoid foramen
The scapula resembles
and receives numerous pneumatic apertures.
in the general
breadth
Its
rests within the coracoid groove.
and there
large
young Casuanus the pro-coracoid
shorter than in Dromanis.
is
that of Dromit'us
in the size of the acromion.
is
long and comparatively slender, tapering rapidly from
The
sternal border forwards to the supra-coracoid foramen.
pro-coracoid
is
There
generally a supra-ctjracoid foramen, but this never receives pneumatic apertures.
whole
is
a large
The
This does not seem to be represented
pneumatic aperture.
any other Palceofjnatha' except, and feebly, some
in
is
At the base of the coracoid and on
sternal border rests in the coracoid grooves.
dorsal surface
its
ligamentous,
leaving a conspicuous notch on the mesial border of the macerated coracoid.
its
large
Cri/pturi.
The scapula is peculiar, curving from the coracoid first inwards then backwards.
The glenoid surface for the humerus lies at the distal end of the coracoid instead of at
its
antero-lateral angle.
In tStnUhio the
size of the pre-coracoid is
and fuse with the internal angle of the base of
finger-like process ultimately to reach
the coracoid, enclosing in this
of the coracoid
is
way
of great width.
huge
a
enormous, extending backwards as a long
The
mention here that the pre-coracoid of
The
fenestra.
sternal (epi-coracoid) border
supra-coracoid foramen
S.
is
We
wanting.
molybdophancs appears
to
might
be peculiar in
its
greater relative slenderness.
In Apterijx the pre-coracoid seems
it
would seem,
at all stages of its growth.
the adult by ligament only
is
to be
;
wanting only
in
A. oweni, and in this species,
In the remaining species
represented in
so that in the macerated skeleton the pre-coracoid fenestra
represented only by the deep notch on the mesial border of the coracoid.
skeleton of A. anstralis mantelli this notch has been nearly filled
extension of bone in the coracoid of the
minute or absent
in all but
side.
left
A. oweni, in which
it
the absence of the pro-coracoid in the latter and
is
it is
supra-coracoid fenestra
is
This point, remembering
large.
is
its
The
In a
up by a secondary
presence in the remaining species,
interesting.
In the Linoniitldda' the coracoid has reached a state of extreme reduction, being
rod-like
and merging almost insensibly with the scapula.
In the jEpyornithidce, in the form of the pro-coracoid,
Rhea and iJromaus.
The coracoid of the
(/nat/ue
—
Cryj)turi diflTers
chiefly in that
it
from that of
its
it
is
congeners
has a well-developed acro-coracoid.
intermediate between
— the
There
fliglitless
is
Pahto-
a well-marked
2
1
2
ME. W.
224
external lateral process at
as in the
young of the
The
is
furcula
PTCEAFT ON THE MORPHOLOGY AND
p.
its
The
members
base.
flightless
pre-coracoid
is
The
wanting.
scapula
is free,
of this group.
slender and U-shaped.
The Development of the Sternum and Pectoral
a.
x\rch.
Sternum.
Miss Lindsay's [48] researches have shown that during development the sternum
undergoes a process of shortening both anteriorly and posteriorly, at least in so far as
its
costal elements
are concerned
the posterior shortening
;
is,
however, more
tliau
compensated by the addition of a long metasternum. That the shortening takes place
both anteriorly and posteriorly is proved by the existence of ribs, which in the embryo
are connected with the sternum, but
The cause
atrophied.
which
of the shortening
of the shoulder-girdle which
is
in the adult
is
become divided therefrom and
" consequent on that posterior translation
at once expressed
by the lengthening of the neck and
the shortening of the trunk in the avian as compared with the reptilian type."
sternal abbreviation
is
common
to both
This
Paheo- and Neognatha\
In Struthio " the 7 days' embryo shows an anterior part which may probably be
compared to the manubrium sterni of mammals." A study of a series of embryos shows
that the anterior lateral processes are an outgrowth of the costal sternum
gnathw these processes are
costal
The
to
;
in the
Neo-
be regarded as vestiges of a former extension of the
sternum anteriorly.
posterior lateral processes in the very early chick, according to Miss Lindsay,
" consist of the ends of the primitive costal bands, preserved and prolonged for the
attachment of the lateral part of the rectus, and subsequently modified ... to afford
attachment to the obliquus externus."
These processes, as existing in the adult, may be considered a part of the metasternum, since the primitive part bears but a small proportion to the later additions.
The
metasteriuiin, both in Struthio
throughout
relatively
life.
somewhat
Posterior
lateral
The
cartilaginous
and
Ithea,
remains for the most cartilaginous
metasternum of
and
Dromoius
Casuarius
is
smaller.
processes
are
wanting
in
the
sterna of Bhea,
Casuarius, and
JJrommus.
each half of the
In Apteryx, according to Parker [7 1], " it appears certain that
sternum is not formed by the antero-posterior union of the whole of the sternal ribs."
.
.
.
In one stage of the series which he examined the sternum apparently increased in
length from before backwards to the level of the fourth sternal
mesiad to meet
rib,
which extended
it.
The metasternum
did
not appear until after the posterior lateral processes had
attained nearly their full length.
In one specimen the metasternum was double, and
PHYLOGENY OF THE PAL^OGNATH.E AND NEOGNATH.^.
in another
it
225
was perforated by an oval foramen, which Parker suggests indicates a
paired origin.
No
adults
traces of a keel
were observed, though such are undoubtedly present in some
4 d).
(fig.
Ossification of the sternum, according to
embryonic
Gadow, begins during the
week of
last
life.
In Struthio and Apteryx a pair of pleurostea only are represented
;
in
Rhea, Casuarius,
and Dro7nmis there are a pair of additional centres representing pro-ostea.
Although there
is
is
no
positive trace of a keel in the developing " Hatite " sternum,
it
generally assumed that such a structure was present in more remote ancestors, and
that
its loss is
the result of degeneration accompanied by decaying, and terminating in
absolute loss of the power of flight.
It is possible,
The
not possess a carinate sternum.
represent this ancient type.
Pterodactyle, and
is
however, that the earliest birds did
Tlie keel was never largely developed
not always present
b.
In Struthio and Apteryx only
is
may
Ratite sterna of the Palwognatlid'
among
well
in the sterna of the
the Chiroptera.
Pectoral Arch.
there a well-developed |)re-coracoid.
Apteryx, however, presents, as Parker [72] points out, " the remarkable circumstance
that in one species oi Apteryx {A. oweni) the coracoid is solid, presenting no coracoid
fenestra and therefore no procoracoid, while in another species
prncoracoid
is
present at a comparatively early age, and
{A. australis)
the
frequently retained in the
In other words, the differences between the shoulder-
form of a ligament in the adult.
girdle in
is
two species of Apteryx are of precisely the same nature
as those distinguishing
the Stritthioiies from the other Ratitse."
It
may even
of the coracoid rather than the
homlogue
In Dro7nceus and Casuarius and in Phea the pre-coracoid
be, as Lindsay has suggested, a
new growth
is vestigial.
of the pre-coracoid of Struthio.
Clavicles occur only in the adult
Promceus and embryo Casuarius.
Pelvic Arch.
The
pelvic arch of the Palceognathce differs from that of the XcognatJue chiefly in
two points
fig.
5)
;
difficulty
:
— (1)
and
The
(2)
whatever
great heiglit of the neural spines of the synsacrum (PL
the persistent ilio-ischiadic fissure.
in distinguishing the Struthious
But though there can be no
from the Neoguathine
these cannot apparently be defined in mutually exclusive terms.
prehensive definition that will cut off the one from the other.
arch
of the
Struthious
shall not be subject to
birds
:
it
difficult
is
any exceptions
;
for
to find
XLIV.
a
There
To keep
definition for
pelvis, yet
is
no com-
to the pelvic
this
which
although the persistent ischiadic fissure
is
ME. W.
226
P.
PYCKAFT ON THE MOEPHOLOGY AISD
undoubtedly a Struthious, because a primitive, character,
Cassowary contradict
yet
the
Rhea aud the
this.
The long and narrow synsacrum, with
the synsacrum of the Pygopodes
;
and
its
this
high neural spines, finds
is
its
counterpart in
a point fraught with some significance to
group, inasmuch as we have already evidence to show that the Pygopodes are
regarded as a piece
a primiti\ e type, and it would seem that this last character is to be
this last
of additioual evidence to this end.
The synsacrum, however, of the Pygopodes differs from that of any Struthious bird
elements
in (1) its much greater lateral compression (2) the absence of any pleurosteal
;
caudad of the last thoracic vertebra
:
aud the exposure of the whole of the fused neural
In Podiciphlidw the post-acetabular ilia
whilst in Coly7nbkhi^ they do not rise beyond the level
between the pre-acetabular ilium.
crest lying
meet dorsad of the neural
of the base
crest,
of this crest.
In the Pygopodes, again, the
iscliiadic fissure of the post-acetabular
ilium
is
reduced
pre-acetabular
to a comparatively small foramen, and the post- greatly exceed the
ilia
in length.
In the elongated synsacrum, the vertical ilia, and the comparatively high neural
spines of the synsacrum the Podicipides resemble the flightless PalceognatJue.
It
may
which
is
be, tlae long,
narrow synsacrum
characteristic only of the
is
primitive,
and that
this
form of synsacrum,
Pygopodes (including Hesperornis) amongst the
Neoqnathw, was derived from a primitive ancestor which had not yet adopted the
more usual Neognathine form of pelvis. The fact that the much more recent and
highly specialized Auks, which have adopted the same mode of life as the ancient
Divers, have a distinctly Neognathine
acetabular
gnathce
is
post-
supports the view that the synsacrum of the P'ygopodes and P'alceo-
primitive in type.
In the
position,
ilia
type of pelvis with widely separated
fli"^htless
members of
and always meets
its
the Palaognathce the pre-ilium
fellow
in the
middle line
is
always vertical in
above
the
synsacrum.
Pleurosteal elements representing the sacral vertebrse are always present.
The
uiEpyornithidce
and Linornithidce, whilst they agree with the remaining Pakeo-
gnathce in the great height of the synsacral neural spine, differ, as
we
shall see,
in.
one or two important features from the other Palceogiiathce, though these structures
are readily traceable back to the more primitive form.
In Prommi.s the pre- and post-acetabular
acetabular
is
markedly the longest.
The
ilia
are subequal in length, or the post-
pre-acetabular ilium has a strongly arched
which caudad sends down u long
the same time as the anterior boundary
dorsal border, a sigmoidally curved ventral border,
acetabular process to join the pubis and serves at
of the acetabulum.
middle.
the dorsal to the ventral border
is
considerable,
innominale bone forming a vertical blade veiy slightl} hollowed in
The post-ilium is narrow cephalad and tapers rapidly from before back-
this portion of the
its
The depth from
PHTLOGENY OF THE PAL^OGNATH.E AND NEOGNATIT.E.
wards, this region of
specimens
it
tlie
pelvis in
quite pre-candad
is
tliis
respect resembling Sfruthio.
in adult birds it
;
227
In half-grown
becomes fused with the extreme
postero-dorsal angle of the ischium.
The ilium
increases markedly in breadth during
its
This increased breadth
growth.
accompanies a corresponding increase in the height of the neural spines of the sacral
and post-synsacral
vertebrie, which, as has already been described (p. 208), in the
embryo are represented only by low cartilaginous transverse ridges. In the embryo the
pre- and post-acctabular ilia are slightly convex superficially, in the adult they
become
slightly conca\'e.
The
It
is
region,
more
or less, of the sacral vertebrae.
immediately above the anti-trochauter, but does not project outwards as in
One
Fliea.
marks the
supra-trochanteric process
lies
of the most remarkable features of the pelvis of Droma'ns and Casuarius
the close embrace with which the post-ilia receive the post-sacral vertebrae of the
synsacrum, almost entirely suppressing the para- and diapophysial processes.
In Dromcexis and (.'amarlus the sacral vertebras extend outwards beneath the vertical
border of the ilium to articulate with the ascending process of the iscliium at
its
junction with the ilium.
The
from that of D. novK-holIandio'
pelvis of I), ater differs
development of the pectineal process
and
is
but slight
— wherein
in that the obturator fissure is wider, the rotation
The
less.
The
crest of the pre-ilium
pelvis of
it
chiefly in that the
approaches Casuarius,
—
backwards of the pelvis being
less arched.
is
Dronueus can scarcely be distinguished from
tliat
of Casuarins.
In the young bird (half-grown) the neural spines of the sacral region of the syn-
sacrum appear
to
be more transversely expanded than in Casuarms.
In the adult the
pre-ilium appears to be relatively of greater breadth in Casuarius and the post-ilium
and narrower.
relatively longer
The
and the width across from one side
strong «^-shaped
is
supra-trochanteric processes
to the other are apparently rarely attained
by the
Cassowary.
Judging from the skeletons
width
across
pelvis of
in
the
Museum
collections
it
would seem that
in
the
the supra-trochanteric process and the relatively long post-ilium, the
Dromceus
is
readily distinguishable from that of Casuarius.
Unfortunately, however, these data do not appear to be really reliable, for in a
jielvis
of C. casuarius the resemblance to these very characteristic points in Bromauis were so
close as to render
In
C. bennetti
it
most
difficult to distinguish
between the two genera.
the width across the supra-trochanteric process
is
as conspicuous as in
Uromnus, but the whole pelvis is of course much smaller than in Dronueus.
In the young (half-grown), both of Casuarius and iJromaus, the ischiadic
complete.
Later in
life it is
fissure
is
closed by fusion of the extreme antero-dorsal angle of the
In very old birds the width of this
end of the ischium fusing with the ilium.
fissure caudad becomes seriously encroached upon by the downgrowth from the postfore
MR. W.
228
PTCEAFT ON THE MOEPHOLOGT AND
P.
ilium of an exceedingly thin plate of bone with a jagged free edge.
region of the 5th post-sacral vertebra, and
is
It arises in the-
From
continued backwards from this point.
the superior border of the ischium there arises a similar plate of bone of corresponding
In very old birds these two jagged-edged plates nearly meet, foreshadowing
length.
the formation of the ilio-ischiadic foramen of the Ncognatluv.
The
The
interobturator process, botli in Casuariiis and
Dromons and
both in
ischia,
Casuarius, are
very distinctly bent inwards towards the middle
;
more
-shaped.
Its
vertical process
it
articulates with
pubis taking place
articulates with
casiLarius,
Cas%(arms
earlier than is the case
straight
(in the
roughly
Antero-
anti-trochanter.
with the vertical process and the ilium.
only slightly curved in
or
the remaining species of
in the living bird
free
this,
posteriorly in
reaching nearly to
the dried skeleton,
with the ischia.
pectineal process seems to be formed for the most part by the descending process
It reaches its greatest
of the ilium.
pelvis of Casuarius casuarius
of Casuarius
The
is
young) and eventually
with a strong outward curve in Dronueiis and
They are always
the ischia.
of
connected by cartilage
The
significance
The ischium
In the adult they extend beyond
end of the post-ilium.
end
rod-shaped and
In the nestling they do not extend backwards as far as the level of the
Castiarkis.
the
less
and eventually fuses with the pubis, the fusion with the
are very slender relatively,
The pubes
free
much
well marked.
this is a point of great
fuses with the pre-ilium, forming the lower half of the
ventrally
is
or
considered in connection with the ischium of lihea.
when
1
Lrommus,
it is
it is
development apparently in Bromwus, hut
only slightly less in
size.
but feebly developed.
outline of the pre-ilium differs, but the series of adult skeletons in the
collection
is
in a
In the remaining species
not sufficiently large to enable any reliable
data to
be
Museum
drawn from
comparisons.
Sjiecific differences in
I
think
it
the form of the pelvis of Casuarius
extremely doubtful.
subject of this paper
is
The
may
possibly be found, but
collection of adult skeletons
which forms the
not nearly large enough to afford the material necessary for
such a task.
The
pelves of the C. henneiti group seem distinguishable from the C. casuarius group
We have
by reason of the relatively longer and narrower post-acetabular ilium.
no adult skeletons of the C. unappendiculatus group.
In Rhea the form of the pelvis is unique, but, as will be shown, is probably a direct
modification of the Uromaeine type.
In the nestling the pre- and post-acetabular
is
abruptly truncated caudad, and rests by
approximated
ischia,
to
its
ilia
are subequal in length.
postero-ventral angle
be described presently.
The
The
upon the
latter
closely
supra-trochanteric process
is
but feebly developed, and the anti-trt)chauter is but slightly below the level of tiie
The post-acetabular ilium is long and narrow and
neural crest of the synsacrum.
—
.
PHTLOGENT OF THE I'ALiEOGNATH^ AND NEOGNATH^.
closely resembles, at this
stage,
tlic
pennanieut condition which
229
bone obtains
this
in StrutJiio.
The extreme
antero-ventral angle of the pre-ilium
outwards, overlapping the penultimate thoracic
The
completely fused.
cephalad and caudad, but leave the neural spines of
ischia,
which,
ultimately becomes
the synsacrum,
synsacrum exposed at the
tlie
sacral region.
will be
it
it
innoniinatcs meet in the middle line above
bottom of a groove over the
The
sharply defined and produced
is
with which
rib,
remembered,
in
Casuarius approached one another in
the mid-ventral line, in Bliea actually meet at a point corresponding with a line passing
tlownwards from behind
ischia
life
more or
The
and a
backwards they remain
These approximated
Tlius a kind of false roof
is
formed
far
beyond the
to the
abdominal
floor to the renal fossa.
pubes, as in Dromaiifi and to a lesser extent in Casuarins, are bowed outwardly
and terminate
The
this point
completely fusing.
less
form a Icng backwaidly projecting median bar, produced caudad
level of tlie post-acetabular ilium.
cavity
From
anti-trochanter.
tlic
inclose juxtaposition, later in
freely near the
pectineal process
end
formed
is
of the ischia.
in part
by the descending acetabular process of the
ilium and partly by the pubis.
The adult
wherein
The
the
the
it is
many
pelvis differs in
respects from that of the nestling, carrying the points
peculiar a stage further in development.
now forms a large overhanging ledge, directly above
acetabulum. The innominate bones now meet in the mid-dorsal line throughout
whole of their length. The post-acetabular ilium fuses on either side by a broad
supra-trochanteric process
expanded foot with the
nestling
—
is
and hejond this fusion
ischia,
continued backwards a long pointed process.
caudad and mesiad, fuse eventually with the
The
—a
point of contact only in the
The
pubes, turning suddenly
ischia.
sacral vertebrae in Casuarius hiy level with
tlu_'
ventral border of the innominate,
and extended outwards to articulate with the
vertical acetabular process of the ischium.
In Rhea these vertebrae are raised up so as to
lie
The
above the level of the acetabulum.
post-acetabular ilium, in the nestling Grebe,
acetabula, in the adult
it is
longer thau
is
subequal
in
length with the pre-
this.
In Siruthio the pelvis presents one or two points wherein
it
differs
markedly from
that of the other Palwognathoe
Like iJronueus, the post-acetabular
difference
is
much
greater in Struthiu.
is
longer than the pre-acetabular ilium, only the
The
and widely separate from the ischium, and
to be almost rod-shaped.
po&t-acetabular region ren)aius permanently
is
so
narrow
The pre-acetabuhir ilium
is
in
proportion to
its
relatively as long as in
rather convex than concave.
length as
Brommas.
The fovea
iliaca anterior
peculiar:
since the pre-acetabular ilium instead of being concave externally, so as to,
on
this account,
VOL.
more or
XV.— P.\KT
v.
is
less
No. IG.
In this particular IStruthio
is
completely embrace the neural spines of the synsacrum,
Deveniber, 11)00.
2 K
ME. W.
230
is
P.
convex, and forms, with
PYCEAFT ON THE MOEPHOLOGY AND
its
fellow of the opposite side, a long
tunnel divided into
right and left halves by the neural spines of the synsacrum, which acts as a
median
partition.
dorsal plane, which in JDrommis, Casiiarivs, and
The
Bhea was
restricted to a small
area between the supra-trochanteric processes, here, in StnitJiio, extends backwards to
Furthermore,
the extreme end of the post-acetabular ilium, though it is very narrow.
downwards on
this plane slopes gently
either side, so that
extremest outer angles
its
approach the anti-trochanter.
pre-acetabular ilium meets in the mid-dorsal line above the synsacral spines the
post-ace tabular ilia remain widely separated one from another throughout the greater
The
:
part of
tlieir
length
old individuals they meet and
in
;
Down
synsacrum.
sacral vertebras of the
fuse with the last two post-
the median space, between the two post-
runs a long bony bar, formed by the fusion of anterior and posterior ossifications
In DinornifhkJa>, it will be remembered, it was pointed out [i]
of the neural spines.
the equivalent ossifications extended laterally, so as to reach the ilia on either side, and
ilia,
thus close what would otherwise form a deep lateral trench lying on either side of the
synsacrum. Tliis trench in Strutliio remains permanently open.
The iscMinn is very slender and rod-shaped. At its posterior extremity it sends
downwards a broad plate of bone which fuses with the pubis. In the young bird, it
The post-acetabular ilium and
should be mentioned, the ischium is free posteriorly.
the ischium remain widely separated throughout
The jmhis
is
life.
In the adult
long, slender, and rod-shaped.
it
extends caudad far
beyond the post-ilium and ischium, and curving sharply downwards and forwards, meets
its fellow of the opposite side in the middle line, and becomes permanently fused
therewith in a long, forwardly directed symphysis.
This fusion of the pubes posteriorly
occurs in no other living bird.
As
is
plate,
well
known now,
attached
skeletons this
is
to
the
the pubis of StrutMo
middle of
generally lost
;
its
further remarkable for a small bony
is
ventral
border by cartilage.
in this case its position
is
In macerated
indicated by a roughened
surface.
This bone appears to have been originally described by Meckel [56], who held it to
More than forty years later this bone
represent the marsupial bone of the Marsupials.
was rediscovered and described by Garrod and Frank Darwin [28]. They also regarded
it as probably homologous with the marsupial bone of the Kangaroo and its allies.
With this view Beddard [7] seems
The pectineal process is long,
also to agree.
slender,
and curved,
its
free
end being directed
upwards.
The
ilio-ischiadic fissure
lJrom(eus and in
Rhea
it
remains permanently open in Struthio.
closes later in
life.
The
obturator fissure
In Casiiarius and
is
closed,
by fusion
PHTLOGENY OF THE PAL^.OGNATHiE AND NEOGNATH^.
of the free end of the pubis with the ischium, both
in
Mea
and
231
and only
Str/ifhio,
in
these.
The
of Apteri/.r resembles that of
pelvic girdle
This
respects.
what we should expect,
is
since
Dromrem and Gasuarim in some
Brommis probably represents the
fjround form from which the other J^nhiorinatJui' have been derived.
In the extraordinary relative length
alone.
It
of the pre-acetabular ilium
extends forward so as to overla]) some four thoracic
acetabular ilium
is
as short relatively as the pre-acetabular
half the length of the pre-ilium.
separated, but the separation
is
is
long
Apferyx stands
The
ribs.
;
post-
never exceeds
it
As in Struthio, the post-ilium and ischium are widely
more marked since the backward rotation of the
is somewhat less than in Sfri/fhio.
The innominates meet in the mid-dorsal
ischium
whereas
in
line as in
Rhea, but with this difference:
Bhea although the median borders meet, yet
widening out and flattening of that
a dorsal plane
is
present as a
])ortion of the dorsal aspect of the pelvis
between the supra-trochanterio processes,
in
Apteryx these dorso-niedian
which
edo-es
lies
meet
along their whole length, save in the legion of the sacrum proper, where a narrow
chink reveals the presence of some three neural spines. There is no supra-trochanteric
process, but
fovea
the
ilia
iliaca anterior
meet
in
the middle line to form a long knife-like ridge.
The
extends backwards into the post-acetabular ilium, terminating
Hehind the acetabulum the vertebral column is bent
shnrply downwards, and has undergone great lateral compression, so that the centra
of the vertebrae have become vertically compressed plates but little thicker than their
caudad of the anti-trochanter.
Furthermore, owing to the downward flexure of the posterior synsacral
vertebrse the post-acetabular ilia have come to embrace the neural spines of the
neural spines.
The synsacrum
extends, moreover, by a single vertebra
of
end
the
beyond the level of the free (caudad)
post-acetabular ilium.
vertebrse instead of the centra.
The ischium
is
a relatively long flattened blade, forming a wider angle with the
ilium than in any other of the Untitw.
The pubis is slender, with a more or less well-marked ventral curve.
Caudad, its
angle
of
postero-ventral
the
ischium, but it never fuses
free end may touch the
therewith.
The
The
obturator foramen and fissure are confluent.
pectineal
process
is
large and pointed, and formed in part by the descendin"-
process of the ilium and in part by the pubis.
Specific differences in the pelves of Apteri/.v are, as
slight.
According to him
A. australis.
to
be the case.
species,
the pectineal process
In our specimens
I
if
there
is
any
is
Beddard has already shown, very
shorter in
A.
oweni than
real difference the reverse
have managed to find
though with
difficulty.
in
would appear
sufficient differences to construct a key for the
Mr. Beddard has already shown that the width across
2 K 2
ME. W.
232
PYCRAFT ON THE MORPHOLOGY AND
p.
the pre-ilium varies, but I think that
characters
may be
expansion of
combined with other
relied on.
Apter)jx anstra^s seems to stand
due
ilium, a width
spite of variation,
this, in
alone in the great width of the pre-acetabular
highly arched dorsal border and a very considerable lateral
to a
A
antero-ventral border.
its
sinuous post-acetabular
rather
ventral
border and a sudden widening of the post-acetabular ilium caudad are combinations
In A. haasfii the pre-ilia are sharply truncated
apparently peculiar to this species.
forwards, and the pre-ilium
is
generally broader in jiroportion than in A. oioeni.
Tlie
post-acetabular ventral border appears to be concave.
A. australis mantelli appears, superficially, not to be very readily distinguishable
from A. oweni.
chief differences appear to lie in the smaller pectineal process and
'Y\\e
broader ischia of A. australis mantelli.
The
of the jEjJi/or/iithida' very closely resemble one
pelves of Dinornithidce and
another, and differ from
post-acetabular region of the pelvis
as broad as long.
This
members
other flightless
all
is
Palwognatim
in tliat
the
flattened out into a large, pentagonal plate, nearly
made up
is
of the
partly by the great length of the transverse
processes of the synsacral vertebr2e, and partly by the great widening of the dorsal plane
of the
ilia
—a
widening only feebly represented among the PaUfoynatha' elsewhere
in StrxtMo.
In the relations of the ischium and pubis the two pelves now under discussion most
nearly resemble Apteryx.
genus the obturator
the foramen
shut
is
Apteryx, however,
ofi"
difteis
one respect, in that in
in
and foramen are confluent.
fissure
from the
fissure.
The
this
In the Dinornithine pelvis
pectineal process
is
large in Apteryx,
very small and wanting in the Dinornithine pelvis.
In the Dinornithidce the sacral are more or
In the yEpyornithidce this
sacral.
is
less easily distinguishable
In the Dinornitliidce the post-sacral neural spines
the floor of which
fossa
is
is
or less completely
plate,
:
middle of a deep
a pair of lateral
from behind forwards
;
slits
fossa,
In the adult this
by a bony roof formed by tabular lateral
e.xpansions from the crest of the neural spines.
quite complete
in the
lie
formed by the upwardly directed neural spines
more
closed
from the post-
not the case.
Caudad, however,
this closure
is
not
run up on either side of the median neural
the extent of these
slits
decreasing with age, but never
entirely disappearing.
In j^pyornis the roofing of the
fossa
is
ample
:
a double
row of foramina only
excepted, which run from behind forwards to the crista transversa.
The
pelvis,
proportion to
both in Dinornithidw and JEpyornithidfe,
its
width than in the other Paheognathce.
is
relatively
much
shorter in
PHTLOGENY OF THE PAL.W)GNATH.K AND NEOGNATH.^i,
The Pectoral
The
pectoral limb
Limb.
the flightless Paheognatlue a very degenerate structure.
in all
is
Perhaps that of RJiea should be regarded as the
In the Wica
The length of
the ante-bracliium.
II.
metacarpal.
similar vestige
manus
The length of the manus is
the jjollex with
In a nestling Bliea
by
phalanv of the second digit
The ante-brachium
is
broad and
—
claw
it;>
And on the
I
has been already detected
the typical Neognathine form
possesses
least degenerate.
length of the extended ante-brachium and
tlie
slightly greater than that of the brachium.
the
233
greater than
in. digit a vestige of a
Wray
in the
the 2nd phalanx
claw
is
The
many long-winged
The
post-axial
is
appearance of the forearm resembles that of
superficial
Neoynathce.
In the humerus the pectoral crest
is
a
pointed.
border of the ulna from the olecranon outwards for some considerable distance
greatly compressed.
;
The idnare
Ostrich.
mere nodule. The 1st
about one-third shorter than the brachium.
is
only very
very nearly as great as
is
in tStriithio this is a
flat,
is
no subtrochanteric
fossa,
is
and the
obsolete, as also
fossa
for
is
There
the crista inferior.
the brachialis internus
only
is
flxintly
indicated.
In Struthio the
less
little
leiigtli
more than two-thirds
In Struthio the manus
In Rhea the forearm
The
the humerus.
in
of the extended ante-brachium and
than the length of the humerus, and
Rhea
it
Rhea the
is
about one-third
short of that of Rhea, being but a
as long as the brachium.
is
longer than the forearm, in
lilica
the reverse
is
about two-thirds and in Struthio about one-third the
distal carpal
mass only just reaches the base of the
is
ii.
digit
more
distal
is
relatively longer in 6'^r«^A?'o than in Rhea.
or less rod-shaped, that of the
end of metacarpal
ii.
is
is
iii.
the case.
leugtli of
1st metacarpal;
extends pre-axiad so as to support the whole base of the pollex.
phalanx of the
in Struthio
is
falls far
manus
The
ii
The
metacarpal
rod-shaped and bowed.
flabellate, the in.
metacarpal
is
1st
In
very slender,
rod-shaped, and sigmoidally curved.
The
much expanded distally by a more or less triangular outits pre-axial border.
The olecranon is but feebly developed. The distal
radius is grooved and much expanded in the direction of its articular surface
ulna, in Struthio,
growth of
end of the
is
for the radiale.
The
pectoral crest of the
There
low, swollen ridge.
humerus
is
is
more conspicuous than
it
receives
Rhea, forming a long,
a wide and shallow fossa distad of the caput humeri,
possibly representing the sulcus transversus.
The crida
in
There
is
also a shallow iucisura capitis.
The subtrochanteric fossa is present but shallow, but
no pneumatic apertures.
The tuberculum medius is moderately well
inferior
is
Manting.
developed.
The
linea aspera for the triceps
is
raised into a long
and sharp ridge, which reaches
MR. W.
234
F.
PYCKAiT
THE MOEPHOLOGY AND
O^'
There
greatest development rather below the middle of the humerus.
its
is
a feebly
developed ectepicondylar process.
The
vadiale
well-developed
is
the uliiare
;
is
reduced
to a small nodule.
In Bromceus, Casuarim, and Apteryx the wing has undergone extreme reduction.
The
In Drommis the length of the whole wing is scarcely longer than the skull.
length of the extended forearm and hand
The manus
about equal
is
to the length of the
There
considerably shorter than the forearm.
is
carpal region, and the radius
is
is
humerus.
no distinguishable
anchylosed with the fused carpo-metacarpal mass.
metacarpals and phalanges have all fused into one indistinguishable rod of bone.
pectoral crest
There
In
wanting.
ever,
it
but faintly represented.
a small subtrochanteric fossa, into which opens several pneumatic foramina.
is
the
is
The
The
skeleton
of
D. ater which
I
had
have
Judging from the measurements given
seems
to
have been
in
examination the wing
for
is
Milne-Edwards's memoir, how-
From the
expected a much greater
smaller relatively than in D. novoe-hollandice.
vestigial character of the coracoid grooves
one would liave
difference.
is
In Casnarins the wing, strangely enough, though relatively smaller than in Droma'iis,
The proportions of the brachium, ante-brachium, and manus
yet less degenerate.
much
are
In the
is
the same as in J}ro7nceus.
adult, the three metacarpals can be plainly
Only one phalanx appears
vestigial.
and
this is
but a vestige.
distinguishable.
— that
of the
ii.
metacarpal,
is
still
The humerus has the pectoral crest
any other Palceognatha\ The subtrochanteric fossa,
to
more strongly developed than in
smaller in Dromams, is very large
In a nestling
be represented
That of the pollex
out.
radiale has anchylosed with the radius, but
The
There appears
to
made
be no ulnare.
in Casuarius.
Casuarius, a radiale,
ulnare,
and
distal
muss
tarsal
are
plainly
distinguishable.
In Ajiteri/x the degeneration of the wing has reached a stage somewhat intermediate
between that of Bromieiis
humerus, however,
it
and Casuarius.
differs
In
the relatively great length of the
from both and resembles Bhea.
brachium and manus are only a
little
metacaipale
in.
In A.
Jiaastii
Parker found an ulnare,
already pointed out, are
radiale,
and a
fairly distinct
In A. austrulis there are no distinct carpals, but traces are dis-
tinguishable of three metacarpals.
The manus appears
The index has two
or three phalanges, the last
In A. oweni the radiale appears to be always distinguishable.
terminating in a claw.
in
ante-
more than half the length of the humerus.
The radius is shorter than the ulna.
The carpus and manus, as both Parker and Beddard have
extremely variable.
The extended
to vary most, according to
Beddard,
in the
matter of degeneracy,
A. australis mantelli.
In an embryo of Apteryx australis, T.
J.
Parker found, not only a separate radiale.
PHTLOGENT OF THE PAL.EOGXATH.E AND NEOGNATH.E.
235
ulnare, and distal carpal mass, but also an intermedium, an element which has so far
been recorded previously
Opisthocomus and Dendroecia.
in
Apteryx there
fact that in the adult
The
as has been pointed out by Parker,
wing of Apteryx
orMn
is
is
a distinct alar membrane, or patagium,
is
strong evidence in favour of the view that the
a degenerate structure, once capable of performing the duties of an
of flight.
A
In jEpyorvis, ajjparently the wing was represented only by the luimerus.
believed to be that of an JEpyoriiis, has been described by Andrews.
to
Its length
was
oi Aptornis, the lower that of
The upper end resembled that of the humerus
Casuarins, "but still further reduced, the radial and ulnar
03
humerus,
mm.
trochlese uniting completely
form a single articular surface."
The Pelvic
The differences between the
many nor great.
In Bromaus the femiathat
is
it
Limb.
pelvic limbs of the existing
differs
from
all
Falwognatim are neither
the other FaUvognatlxi., save Strnthio, in
highly pneumatic, and bears on the ventral aspect of the femur a large
pneumatic foramen.
The popliteal fossa
The til'w-tarsus
between the great trochanter and the head of the femur.
It lies
is
is
of moderate depth.
not easily distinguishable from that
difference appears to rest in
This vertical direction
tlie
slighter
rather marked
is
and
less vertically
The
chief
extended procnemial
crest.
of Casuarius.
Yhejibula extends to the lower
in Casuarius.
fourth of the shaft of the tibio-tarsus.
The
tarso-metatarsHs
as long
is
grooved in front, and has a
as the tibio-tarsus,
moderately developed median keel to the hypotarsus.
The proximal phalanx
the proximal phalanx
The
is
of digit
long, the
is
relatively long, the
2nd and
anterior metatarsal groove
toe, in
each
is,
The tarso-metatarsus
Tlius
it
The
is
is
non-pneumatic.
is
much
The
has lost one phalanx.
position of the pneumatic fossa in
The
The
tibio-tarsus, as already pointed
shorter than the tibio-tarsus,
pierced by a foramen dorsad
of
is
very deeply grooved
the scar for the tibialis anticus,
can readily be distinguished from the tarso-metatarsus of Dromceus.
ungual phalanx of the inner toe is greatly elongated in other respects the size
;
of the phalanges closely agrees with those of
digit
in digit IV.
however, somewhat deeper.
Jjromaus may be indicated by a minute aperture.
out, differs but slightly from that of Vronucus.
and
;
especially the 3rd and 4th are very short.
foot, in this skeleton
In Casuarius the femur
anteriorly,
2nd very short
of D. ater closely resembles that of D. novw-hollandice.
pelvic limb
The outer
li.
II.
is
relatively shorter.
Dromaus, but the proximal
plialanx of
ME. W.
23G
PYCKAiT
P.
THE MORFHOLOOr AND
0.\
In Strutliiu the femur, as iu Dromaim,
highly pneumatic, and, as in Dromceus,
is
bears a large pneumatic fossa between the head of the femur and the great trochanter.
The
shaft
condyle
is
much
relatively
is
much
thicker than in Dromceus or Cassowary, and the fibular
larger than the inner tibial condyle.
The groove
them
dividing
is
also
narrower and deeper than in Dromceus.
The
size
Irom
tibio-tarsus, apart
its
greater
and hooked form of the ectocnemial
which extends
is
is
peculiar iu
fibula,
upper border of the proximal
tarsal
Furthermore, the lateral borders of the
shallow.
posterior trochlear surface are produced backwards and
proximal ridges at right angles
The
tarso-metatarsus
It
trochlea.
the relatively smaller
and the great length of the
crest,
to within a short distance of the
The extensor groove
mass.
size,
upwards into
a
of
pair
long axis of the shaft.
to the
of course at once distinguishable by the absence of ento-
is
grooved anteriorly, and bears a pair of foramina dorsad of the scar for
is
the tibialis anticus.
The ungual phalanx of the outer toe is represented by a small nodule.
It may be at once distinguished from
In Rhea the femur is non-pneumatic.
that
of any other Faheognathce by the great depth of the popliteal fossa.
tarso-metatarsus resembles that of Strutliio in the feeble development and in
The
the form of the ectocnemial crest and in the backward projection of the condyle of
its distal
extremity.
The femur
falls far
short of the level of the tarsal mass.
The
tarso-metatarsus has a shallow groove anteriorly, a deep fossa above the scar for the
and a very broad and low intercotylar process. The hypotarsal keel
In Dromceus and Casuarius it is median in
the inner side of the median line.
tibialis anticus,
lies to
position.
The
of the
1st
phalanx of digit
IV. digit
is
ii.
conspicuously lengthened, the 2nd
the proximal phalanx
In AjJteri/x the lemur
long, the 2-4 very
is
and more slender than
relatively large
is
FaheognatJiw, save the Tinamous.
It is
is
very short
short.
in
any other living
non-pneumatic, and the popliteal fossa
is
deep.
The
tibio-tarsus proximally
is
much
ecto- nor procnemial crests are
intercondylar gorge
The
is
tarso-metatarsus
;
The extensor groove
developed.
deep, and the ento-condyle of great
the intercotylar tubercle
anticus
broad and flattened from back to front, neither the
is
is
shorter than the femur,
well
marked
;
there
is
much
is
deep.
The
size.
flattened
from back
to front
a fossa above the scar for the tibialis
the 2nd and 4th trochlese are widely separated from the meso-trochlea.
The
hypotarsus bears a pair of low median ridges.
There are four
digits.
The
digits contain
no conspicuously shortened or elongated
phalanges, and therein difler from the other FaUeognathxe herein described.
In Dinornis the femur
distal
end
is
is
relatively
of great width
;
much
shorter and thicker than in Struthio.
the intercondylar gorge
is
Its
very wide and shallow and
—
PHTLOGEXT OF THE PAL.EOGNATH.E AND NEOGXATILE.
extends proximad on to the shaft, terminating rather behind
The
no conspicuous pneumatic foramina.
trochanter
i^reat
237
There are
middle.
its
very large
is
it is
;
pro-
duced upwards above the head, and not backwards caudad of the head, being an
exaggeration of what obtains in
/h'i»nn'/'s
and
The
Ca-snariiis.
popliteal fossa
is
well
defined, but not deep.
The
much
tibio-tarsus is
fiatteued from front to back, as in
the ectocnemial crest is large,
bone a
flabellate
may not be
form
inflected
the procnemial crest
;
Aptevyx and j^pyornis
;
and gives the anterior view of the proximal end of the
— according to
tlie
There
genus.
The
feebly developed.
is
is
may
shaft
or
a strong extensor bridge and an
intercondylar tubercle.
In jEpyoriii^ the femur
The
produced caudad.
is
The
also very short.
intercondylar gorge
great trochanter
is
very lofty and
wide and deep, and the popliteal fossa
is
shallow.
The
tibio-tarsus has
and large
procnemial
There
inflected.
is
proximal end much flattened from front to back; a feeble
its
directed
laterally
ectocnemial
no extensor bridge, and the groove
the
Distally
crest.
sliaft
is
siiallow.
is
The tarso-metatarsus is short, wide, and grooved anteriorly, but the groove is shallow.
The ectotrochlea is widely se[)arated from the mesotrochlea. There is a low, median,
hypotarsal ridge.
Phalanx
of digit
1
is
li.
moderately long,
digit IV. are conspicuously short, the 4th
is
2nd
tlie
The
shorter.
short
is
phalanges
;
3 of
2,
proportions of the phalangeals
agree closely with those of Lromwiis.
In working out the development of Apterya-, Parker [71] found, in the earlier stages,
all
five digits
present
cartilage attached by
—
as
in
some
'Neorjaathn'.
proximal end
its
The
to the fibulare,
fifth
and by
digit
its
was a short conical
prc-axial border to
tlie
distale.
The
The
was of the same length as the
fibula
tarsus contained
the
usual
tibia.
elements
—a
and
fibulare,
tibiale,
The
distale.
ascending process of the tibiale at no stage showed any sign of a separate origin,
comparable
articular
The
to
Later, after hatching, there appears in the mesotarsal
an intermedium.
pad a pair of
Sometimes only one
centralia.
is
present.
procnemial crest ossifies, as usual, from a separate centre.
would remark that
ossification I
ficance.
Rather
it
would seem
the great trochanter of the
strain.
In a recent paper
unknown
to
me
it is
to be
doubtful whether
comparable
shaft by
VOL. XV.
cartilage,
— PART
No.
V.
is
however, described
as
an epiphysis, as
I,
the late
W.
it
K. Parker
this
[79].
epipliysis.
December, 1900.
In
also,
my own
procnemial ossification
large to form a complete
a true
17.
separate ossification-centre of
indicate a position of great
sufficiently
as
has any phylogenetic signi-
to
at the time, did
—was
to the
this
mammalian femur, and
claim some justification, since the base of
young Grebe
it
Concerning
tibial cap,
Moreover, the
though
case
—
I
may
that of a
divided from the
resemblance
to
2 l
a true
MK.
238
epiphysis
I will
the more complete in that
is
not
PYCRAFT OX THE MORPHOLOGY AND
"W. P.
comment
it
articular surface for the femur.
affords an
further upon the matter here as I have the subject
still
under
investigation.
MYOLOGY.
the Archwopalatime
The musculature of
of Beddard, Fiirbringer,
is
must not be supposed, however, from
On
the contrary,
many
well known, thanks to the researches
Gadow, Garrod, Haughton, Meckel, Owen, Parker, and
So well have these been done, that, gleaning
It
now
this, that
the subject
points yet await investigation
here, however, either because material for this paper
to
be of
little or
no value so
far as
to be
developed dermo-dorsalis.
base of the skull
down
now
add.
quite exhausted.
mention of these
finds
no place
was lacking or the points appeared
Skin-muscles.
most strongly developed
and Dromceus appear to be peculiar
Casxiarius
;
is
little to
was concerned.
it
The
The dermal muscles appear
have but
late in the day, I
others.
in
Apteryx.
in
that
they
possess a strongly
In a Casuarius miappendicidatus this extended from the
the skin of the neck and back along the middle line.
At the
base of the neck the muscle gives place to tendon, which, becoming rapidly stronger,
is
inserted,
by means of numerous long, coarse, tendinous
fibres,
along the back as far
as the pre-acetabular ilium.
The Muscles of the Wing and Shouldek-girdle.
we may attribute the absence of many muscles found in
Furthermore we must regard this loss as one of considerable antiquity,
Neoqnatlue.
since the flightless members of the Neognathce exhibit no such reduction in tlie number
To
of flight
the loss
of the muscles concerned in this form of volition.
Mr. Beddard
the
[7] has given an
They
Palmognathce.
are
admirable summary of the missing wing-muscles of
the pectoralis
propatag talis,
biceps
propatagialis,
deUoides propatagialis, deltoides minor (excluding Apteryx), scapulo-Immeralis anterior,
and expansor secnndariorum.
wing of JRhea
The form
a
(see
fig.
Concerning the
last
mentioned, traces occur in the
5 a, p. 240).
of Jiexor carpi ulnaris of
Rhea
is
extremely interesting, as will be seen by
comparison of the accompanying figures (pp. 240, 241).
It will be noticed that in Rhea (fig. 5 a) this muscle
(1) a pre-axial ribbon-shaped and partly tendinous, which
and
(2)
1st
metacarpo-digital
portion
a post-axial,
is
much
larger,
remex.
The
is
is
divided into two parts
inserted on to
strap-shaped, fleshy portion inserted
post-axial
bounded by a narrow thread of tendon
tlie
:
ulnare,
on to the
border of this hinder strap-shaped
— a rudimentary vinculum
elasticuni.
PHTLOGEXY OF THE PAL.EOfiNATH.E AND NEOOXATII^.
we have
In the Crypturi
a second modification
5
(fig.
240).
b, p.
The
239
jiost-axial fleshy
moiety has here apparently degenerated so as to be represented only by the
vestigial
muscular slips from the body of the pre-axial portion, which is thick and fleshy.
vinculum elasticnm represented
nascent, not vestigial
suppose that this
—
is
in
Bhea by a yiidiment only
The
mean
We may
— by rudimentary
here increased to form a great tendinous
slioet.
I
an improvement on the proto-carinate form seen in lihea.
In Opisthocomus (fig. 5 c, p. 241) both pre- and post-axial moieties are fleshy. The postaxial shows the first indication of the complex vinculum elasticum of the Neognatlm.
In Leptoptih(s the pre-axial portion is almost entirely tendinous. Proximally, the
is
tendon divides, one branch running directly
the ulnare, and one fusino- with the
anterior border of a muscular post-axial portion throughout the greater part of its
to
length, finally trending upwards to join
the main pre-axial tendon on the ulnare.
muscular, post-axial portion apparently represents the broad strap-shaped
portion of Rhea
it differs therefrom only in that it is inserted into the first
cubital
This
last,
;
instead of the
first
metacarpal remex.
There are numerous other modifications, varying
in complexity, to be
found amono-st
the Neognathw, a study of which would probably yield important results.
The wing
facts
Rhea
of
represents, probably, as Dr.
Gadow
—remarks, a proto-carinate form.
In the manus of Rhea, Mr. Beddard writes, "
There
than in Struthio.
some of these are much
"The
are, in
the
first
—
to
whom
we have more evidence
I
showed these
of defeneration
place, only twenty-one muscles at most,
and
simplified.
muscles that appear to be totally wanting nre (1) the extensor digiforum
(2) the pronator profundus.
communis,
"The
extensor indicis
flexor suhlimis
may
is
only represented by the belly arising from the wrist.
possibly be
represented by a slip of muscle arising
The
from
the
tendinous edge of the flexor metacarpi uluaris ....
" The origin of the biceps is peculiar it arises not only from the coracoid spine by
;
a rounded tendon, but also by a sheet of tendon edged with muscle from the whole of
the coracoid and from just an adjacent Int of the sternum.
It is inserted
on
to
both
radius and ulna.
"
As
there
in the Ostrich, the radio-metacarpalis ventralis arises
is
a special peculiarity in the presence
Finally
Of
tlie
ectepicondylo-ulnaris
Struthio, Mr. Beddard writes that, " in spite of the small size of the
musculature.
oi Stnithio
that
to its insertion to the extensor indicis.
distinct."
relatively to that of flying birds, there
its
On
are less
In Rliea
of a muscular slip running from the
tendon of the extensor metacarpi ulnaris near
is
from the ulna.
is
but
little, if
the contrary, indeed, for
it
manus ....
any, evidence of degeneration in
might be
said
that the wing-muscles
degenerate, or at any rate less modified, than those of Curinales, in
amount of muscle compared with tendon
is
greater.
The complication
2l
2
of the
ME. W.
240
P.
PTCEAFT ON THE MOEPHOLOGT AND
Fig. 5.
A.
B.
i>.e.
conjoined flexors digitorum
It
is
highly suggestive of a walking or climbing animal.
seems to be conceivable that the Ostrich branched
off"
from the avian stem before
the power of flight was perfectly established.
"
The
biceps oi Struthio arises
from
tlie siyina
and ulna and the membrane between them.
The
coracoidea and
The
is
inserted on to the radius
extensor metncarpi radialis
is
ectej)icondi/lo-ulnaris is absent or fused with the extensor metaccirjri ulnaris.
extensor digitorum
one muscle.
The
single.
The
communis supplies only the index. The two ^^''"onators form only
fiexor digitorum suhliinis and jjrofundus arise by a single head
from the flexor condyle of the humerus.
upper part (:=sublimis) ends
in
The two muscles immediately
two tendons, of which one
is
inserted
divide; the
on to (the)
PHYLOGE.N'Y OF THE PAL/EOGNATH^
AND NEOGNATH^.
241
fig. 5.
C.
eat.
t^.e
— Inner view of a
A.
wing oi Rhea mneneana,
dissection of the
ariorum and the
? proto-carin;ite
to
show the remains of the expausor sceund-
condition of the flexor eurpi iihiaris.
portion eventually- inserted into the base of the 1st primary
runs along
B.— Inner
its
— Inner view
of
C'alodromas deijans.
carpi idnaria has become reduced
remiges, wliilst the vineulum
C.
large, fleshy inferior
free border, probably representing the i'incuJum elasticinH.
view of a dissection of the wing
the fle.cor
Note the
remex and the thin tendinous band which
elastieuiit
of a dissection of the
Note that the
to slender slips of
fleshy posterior portion of
muscle to the bases of the secondary
has greatly increased in size to form a broad tendinous sheet.
wing of Ojikthocomiis
eristatus,
showing another phase
in the develop-
ment of the vinciiJum elastkum and the persistence of a strongly developed posterior belly to the flexor
carj)! uhiaris.
Numerous intermediate
eap.scc.
f.c.u.
uJn.
li.e.
=
stages occur
amongst the
Neot/natluf.
expansor secuadariorum.
= flexor carpi ulnaris = anterior, " = ))osterior
= ulnare.
= vineulum elasticum.
'
:
portion.
with the upper tendon of profundus, and also gives oflf two
of the
slips which surround that tendon and, reuniting, fuse with the lower tendon
The lower part of the muscle (=profundus) gives oif two tendons, of
profundus.
radiale, the other fuses
which the upper ends on the
last
phalanx of the index.
"
The Jlexor metacarin
this bone, receiving also
first
metacarpal, while the lower runs to the base of the
ulnaris ends fleshily on the ulnare, but
some
fibres
from
it
is
prolonged beyond
to tlie metacarpal.
from the ulna and not from the radius.
" The total number of muscles in the hand of the Ostrich is twenty-three, allowing for
the absent ectejjicomJylo-uhiaris. The additional muscle is a small pronator quadratiis."
"The mdio-metaearpalls
veniralis
....
arises
MR.
242
PTCEAFT ON THE MOEPHOLOGT AND
\V\ p.
In A2)teryx the usual flexors and extensors of the forearm are present, and, as the
Prof. T. J. Parker points out, a rather unusually large development of muscles
late
acting as pronators and supiuatoi's for so small and vestigial an organ.
The
biceps of Apteryx
only on to the radius.
Gallinaceous
the
single-headed, arising only from the coracoid
is
;
inserted
it is
Apteryx differs only from the other genera in the possession of
and Tinamiue entepicondylo-ulnaris and an accessory hrachialis
anticus.
In Casuarius and Drommis the wing seems
to
have reached
most
its
vestigial
condition.
The
hice])s in
Casuarius arises only from the coracoid, and
is
inserted by a single
In Bromceus, according to Beddard, the biceps
tendon both upon the radius and ulna.
has the same peculiar origin to that oi Bhea
(p. 241).
the Struthious birds," writes Beddard [7], " except Apteryx, have lost the
" All
metajjatagialis,
serratus
the
latissimus
and
metapatagiaUs,
dorsi
the
i)ectoralis
abdom'malis.
"
On
the
the other hand, Apteryx has lost what the other Struthious birds have retained,
Jatisshnus
howevei',
" It
is
and the rhomboideus profundus
dorsi anterior
;
the
muscle,
latter
not distinguishable in the Cassowary.
must be admitted,
that Apteryx, so far as concerns the anterior
therefore,
extremity, has diverged from the hypothetical ancestral condition in slightly different
lines
from other Struthioiies."
The rhomboideus
RItea
it is
superjicialis in BItea
and Struthio
inserted on to botli the coracoid and
In
arises as in Neognathce.
scapula, in Struthio on to the scapula
only.
In Apteryx, Casuarius, and
Bromwns
The rhomboideus profundus
Casuarius
it is
with
in
it
arises
from the
ribs.
Casuarius and Broinceus arises from the
from the serratus profundus, and on
difficulty distinguishable
In
ribs.
this
account Fiirbringer [22] inclines to disallow its existence as a separate muscle.
The
serratus superjicialis of Apteryx retains the pars metapagialis, which
is
wanting
in the other Paheognathce.
The
coraco-hrachialis internus of Casuarius
Furthermore,
in this last
genus
it
is
entirely tendinous; in
extends on to the sternum.
Rhea
In Struthio
largely so.
it
is
larger
than in any other PaJceognathoe.
Thigh- and Leg-muscles.
Struthio and Apjteryx only have
formula
the
leg-muscles complete, which
make up
the
AB.XY+.
In the StrutJtiones generally the accessory femoro-caudal (pars iliaca m. caud-iliof'emoralis) calls for special
comment.
In
all
save Apteryx
it
is
characterized by
its
PHYLOGENT OF THE PAL.EOGNATH.^ AND NEOGXATHvE.
great
Strutkio and Casuarms being rightly described as enormous.
size, in
caudal, on the other hand,
is
relatively
Bhea and Dromwus both agree
in
24?.
The femoro-
but feebly developed or absent.
having
lost the
Fis.
femoro-caudal.
fi.
cr
^ / TncMC a/7zi
max pes
^/.
t.
Left side view of a dissecfion of the thigh-muscles of Casuai-ius unajijiendkidatus, to show the accessoiy
The accessory adductor,
Stnithious adductor, the femoro-caudal, and accessory semiteudinosus muscles.
femoro- and accessory femoro-caudal
= the
caud-ischio-ilio-femoralis of
= accessory Struthious adductor.
= biceps.
f.e. = femoro-caudal.
acc.f.c. = accessory femoro-caudal.
Acc.ad.
ijljiiui.
's.t.
=
gluteus minor.
yj.mx.ijost.
=
=
,,
posterior.
(jl.aiU.
=
,,
anterior.
'jl.mx.ant.
h.
=
Gadow.
maximus
„
anterior.
Sat: z= sartorius.
semiteudinosus.
= accessory semiteudinosus.
si/ih.
=
semimembranosus.
adductor longus.
icl.c.
=
ectocnemial crest.
cr.
=
=
crureus.
ad.l.
adductor longus.
gl.med.
=
gluteus medius.
=
=
(tcc.st.
ad.l.
The femoro-caudal muscles
by Garrod and Beddard.
independent dissections.
p.b.
pubis.
of Apteryx are peculiar and are differently determined
Garrod's view
Garrod [30] says
is
:
adopted in the present contribution, after
— " The accessory femoro-caudal
is
peculiar
;
MR. W.
'244
for,
PYCRAFT ON THE MOEPHOLOGT AND
P.
on removing the biceps
cruris,
its
portion
superficial
seen running obliquely
is
upwards and forwards to the whole length of the linea aspera of the femur, from
The sciatic artery and nerve are
Owen) and parallel to its insertion, as
usual origin.
longus of
superficial to
most birds
in
case except in the Struthiones and Criipturi, perforate
it
this
muscle (adductor
but they, as
;
its
is
not the
at the ischiadic notch, leaving
a small portion of the muscle (the adductor brevis of Owen) above them.
The
anterior
terminal fibres of this muscle are situated external or superficial to the accessory semitendinosus.
" After this muscle has been removed or turned back, there
which,
if
is
seen a deeper muscle,
the one described above did not exist, would be justly considered to be the
femoro-caudal and the accessory femoro-caudal, part springing from the
iliac
ridge and
part from the coccyx, whilst both are inserted into the posterior portion of the linea
aspera and have the nerve to the semi-membranosus situated between them and the
adductor magnus."
it
This deeper muscle Garrod does not further distinguish.
Probably
should be regarded as representing the obturator externus.
The femoro-caudal,
memhrano.ms
it
should be remarked,
is
very slender
In Garrod's paper he appears to associate
.
it lies
;
deep of the semi-
it first witli
the superficial
and then with the deeper muscle.
Mr. Beddard apparently regards the obturator externus of this paper, and the deep
layer of muscle described by Garrod, as tlie muscle which would be justly considered
the accessory femoro-caudal, in the absence of the superficial layer just described.
The muscular mass embracing the emergence
of
the sciatic
nerve and artery
should be regarded, I would suggest, as the Struthious adductor referred to by both
Garrod and Beddard, but fused distad with the gluteus anterior, and caudad with
A
the accessory femoro-caudal.
comparison of
fig.
6, p.
243, will serve to show the
probability of the correctness of this interpretation.
The semitendinosus and
similar.
in the
In Bhea the former
Cassowary
is
means of a long tendon,
inserted on to the tibia by
two former genera
at least of the
In StvKtJiio the accessoiy head
accessory head
Casuarius, Dromceus, and Rliea are very
accessory in
its
is
loses itself in the gastrocnemius.
In Apteryx, according
small.
shows a slight variation
it
in
its
relations
within
to
the
Beddard, the
" In
genus.
A. anstralis the accessory head of the semitendinosus was distinctly separated from,
In A. haasti no such
though parallel to, the middle head of the gastrocnemius.
separation
is
obvious."
The semimembranosus
(ilio-fiexorius,
head arising from the ischium
;
Gadow) of Apteryx
between these two heads
two-headed, the second
is
the head of the femoro-
lies
caudal.
In Lromceus the semimembranosus, according
to
Garrod,
is
peculiar in that
it
has
au aponeurotic connection with the middle of the linea aspera.
In Casuarius
it
is
but feebly developed
;
it
passes rapidly into a thin
flat
tendon to
PHYLOGEXY OF THE PAL-EOGNATII^ AND NEOGNATH,^.
be inserted into the shaft of the
of the gastrocnemius
In Bhea
it
tibia,
below the neck.
border of the accessory semitendinosus.
of the semitendinosus, distad of
passes the middle head
terminating distad of the ventral
its
Its antero-dorsal
accessory head.
angle fuses with the tendon
Its postero-ventral angle
sends
a long slender tendon to the gastrocnemius, to be inserted a short distance
downwards
above the
it
gives off thereto, at right angles, a delicate tendinous band.
it
small, sharply truncated forwards,
is
As
245
tarsal joint.
In Struf/iio, according to Garrod, the semimembranosus tendon fuses with that of
the accessory semitendinosus and then passes down the leg as a long thin tendon to
Thus
the tendon of the gastrocnemius.
The
amhiciis
however,
closely resembling that of
wanting only, among the StrutJtione^, in
is
Mea.
Catmariiis,
Dromieii-'^.
generally regarded as wanting this muscle.
is
In Strut Ii/'o
its
origin
is
the ilium instead of the pectinal
peculiar, being from
process of the pubis.
from the pectineal process of the pubis; as usual, it is
It does not, according to Beddard, give
inserted into the flexor perforatus digit! Ii.
" Furthermore, these
off slips to the other two muscles of the perforatus complex.
In Aptcryx
its
origin
is
small tendinous bands, accompanied by muscle-fibres, and forming a thin, flat sheet of
tissue, are continuous with the ambiens tendon above, and appear to be in connection
at the other
I
end
witli the short
arm of the biceps
sling,
and
to arise
upon this sheet of muscle and tendon as a second
look
from the
fibula.
head of the flexor
question, and as corresponding to the fibular head of birds, such as Ntjcticorax.
this
be
so,
ambiens
same
is
Mitchell's contention that the ligamentous head in question
not so certain as
state of affairs in A.
it
seemed
to be, for
is
both occur in Apteryx.
in
If
a rudimentary
I
f(mnd the
haadi."
In Bhea americaua, Gadow [25] found the ambiens to be typically developed. This
I am able to confirm from my own dissections.
ambiens tendon, at the level of
I would remark that in an adult of this species the
the fibrous tissue of tiie lower
the head of the fibula, sent upwards a tendinous slip to
fl. perf. d. ill. and
border of the tendinous insertion of the crureus and origin of the
tendinous portion of tlie origin of the
perf. et perf. d. 11., and a few fibres to the
common
belly of the
fl.
il.-iv.
and
perf. et perf.
111.
" the muscle arose from the latero-dorsal aspect
In B. inacrorhyncha he found that
pubic spine, and at the same time from the big crural vein, the muscular and
of the
ventral aspect of
aponeurotic fibres of the muscle having firmly got hold of the
The tendon of the muscle passed the knee in the typical way.
Yein
»
In B. (larwini,
usual, but
its
2
tendon,
,
tliis
the m. ambiens of cither side arose from the pubic spine as
before reaching the knee, became flattened out and attached
and very thin aponeurosis to the patella, in a similar
itself with a broad fan-shaped
In a
portion of the m. femoro-tibialis (m. vastus)."
style as the median additional
2 m
yoL. XV.— PART V. No. IS.— December, 1900.
—
246
MR. W.
PTCEAFT ON THE MORPHOLOGY AND
P.
male B. danvini the ambiens of the
thigh stopped at the knee, as in the female,
left
bnt in the right thigli the muscle was typically developed and passed the knee with a
strong tendon.
In Casuarius the ambiens,
Gadow,
Dr.
according
to
Garrod,
Concerning
wanting.
is
this,
ambiens of Rhea, a portion of which we have just
in describing the
quoted, continues, " This abnormal condition of m. ambiens, which seems to be prevailing in
Darwin's Bliea,
ra.
ambiens and such forms
of
its
the
we
in which, as in Casuarius, this
muscle has
lost still
media m. femori
comparison
of
ambiens muscle.
come
of
to the
The assumption
of
vasti.
s.
formation
the
should, with Garrod,
tibialis
Without an elaborate examination
these
muscles
with
their
nerve-supply,
conclusion that Casuarius did not possess an
still
further reduction of the distal portion of
the m. ambiens explains what I have observed in some Passerine birds,
specimen of Lanius
hentef,
in
strongly developed, received in
head from the pubic spine.
the ambiens muscle, which
"
The
e. g.,
more
independence, and then only forms an additional head of the median part of
portio
and
between a typically developed
the intermediate stage
is
which the m. femori
its
case above described
now
is
(].,
in a
internus, besides being
proximal part a thin spindle-shaped semitendinous
This additional
is
tibialis
e.
little
slip
is
probably the last trace of
generally lost by the Passerine birds.
one way in which this muscle gets
Ciconia and Phoenicopterjis, the reduction does not begin by
lost
its
;
in other cases,
tendon becoming
attached to the neighbouring tendons in the knee-region, but the whole muscle shows
a diminution of
e. g.,
in
its
tendon and belly to a mere thread,
till
Abdimia and Xenorhynclms."
The muscle-formulse of the Pakeognctthce
Struthio
is
as follows
:
at last this also disappears,
PHYLOGENY OF THE PAL^OGNATH^ AND NEOGNATH^.
247
NERVOUS SYSTEM.
The
Parker has shown
large size,
compared with the brains of many Neofjnathw
that,
Emu, Goose, Pigeon,
— the
and further
Brain.
cerebral hemispheres
way they overlap the cerebellum,
peculiiir is the
—Turkey,
Apteryx are proportionately of
in
bearing a very close resemblance to the Passeres.
in this respect
This overlapping conceals the
grooves wliich mark the anterior region of the cerebellum, the hemispheres extending
as far
back as the
The
brains.
flocculi.
The
posterior grooves are visible superficially, as in other
pineal peduncle, in consequence of this overlapping,
is
inclined from
the vertical position backwards.
The
position of the thalamencephalou
also peculiar, in that this region
is
tilted
is
backwards, the lamina terminalis looking upwards, the foramen of Monro becoming
postero- instead of antero-dorsal.
TuE Eye.
The
pecten
Dromceus the number of
14-20
developed
well
is
in
all
the
Pal(eognatha\
reduced to four
its folds is
;
In Apteryx the pecten
The pecten
is
generally believed to be wanting in the adult, though
is
present in Jipteryx, though
not fan-shaped, and plicated.
never conical
In
little
of the disc
is
not really the case.
assumes a peculiar form, being columnar,
a vertical, plicated, densely pigmented septum
left
half,
The eye
seen.
it
is
other birds the pecten has two or more plications
all
— usually forming
dividing the disc into a right and
very
In
Dr. G. Lindsay Johnson, by a series of drawings recently
exhibited at a soiree of the Royal Society, has shown that this
is
Apferi/x.
in numbei'.
present in the embryo.
and
save
in titnitJiio the lamellse are from
is
although
its
base expands over
Hence the apex
small.
it
so that
of the pecten
penetrates the vitreous to a distance nearly equal to that of the posterior surface of the
lens from the retina.
direction have related almost exclusively to the
Dr. Johnson's researches in this
mammalian eye
;
and they have been most
strate a precisely similar " pecten
Golden Agouti
mentioned
—
as
soiree in
he showed
June of
""
fruitful
in the
by means
this year
of
him to demonMammalia, that of the
in results, enabling
eyes of several
drawings,
exhibited
at tiie above-
— bearing a most striking resemblance
to
that of
Apteryx.
JaCOBSON's CARTir.AGES.
These, according
lying parallel to,
to
Parker, are represented by a pair of vestigial rods ol cartilage
and either immediately dorsad or
slightly laterad of, the dorsal
2
M
2
edge
ME.
248
of the vomer.
It
PYCRAFT 0\ THE MORPHOLOGY AXD
"W. P.
can he readily prepared in a well-macerated skull by carefully
removing the vomer.
was
It
first
described by
W.
K. Parker [77]
in
Rhea.
THE ALIMENTARY CANAL.
Buccal Cavity.
The buccal
cavity of the Palcfognatlice presents one
from that of the
differs
NcogimtJuv
XLV.
(PI.
noticeable feature wherein
1)
fig.
:
since, in
it
the former, ihe
posterior nares, or choanse, form a wide, open, more or less cordiform aperture, divided
by a median septum into right
choanee are
In
a
all
slit-like.
and
The Eustachian aperture
the Palwognatlicc the tongue
is
moities
lateral
left
in
the Ncognatlm the
immediately caudad of the choanse.
lies
more or
less vestigial.
The
glottis is also
In Casuarms the tongue has a
wider aperture than in the Xeocpiathce.
relatively
;
denticulate free edge.
The
"On
Dr. Gadow, in his paper
the flightless
CoiNvoLUTioNS of the Intestine.
the Intestinal Convolutions in Birds" [26], considered
members of the Palwognathce
a very heterogeneous group, because
of
the great diversity in the length and arrangement of the main gut and in the develop-
ment of the caeca. " In none of these birds has it come to the development of closed
and well-developed loops of the mid-gut (with the exception of the duodenum). In
Their
this respect they represent the lowest type amongst the recent birds
Nearest of them to the latter comes
connections Avith recent Carinataj are distant.
A]jteri/x through more defined loops, and the Crypturi seem to represent the link.
.
having the second
All the Archceopalatince agree with each other in
and the third left-handed
Gallince, Opisthocomus,
Amongst
;
and
the JS^cognatha'
,
this is a feature
looj)
which occurs again only
.
.
right-handed
in the
Crypturi,
in the Cuculidm."
Avill
it
be remembered,
Gadow was
enabled to demonstrate
a greater harmony, the convolutions of the intestines being always referable to one of
seven types.
Dr. Gadow's conclusions,
it
should be remarked, are based upon a study of the
coiling of the intestines within the body-cavity.
Mr. Mitchell
point of view,
vessels
[60],
who approached
viz.,
after its
the study of the intestinal tract from a different
removal from the body, and the severance of certain
and secondary connections, which reveal the nature of the convolutions of the
gut along
its
attacnment to the ventral border of the dorsal mesentery, and the
position of the diverticulum cfecura vitelli and
vein, entirely
confirms Dr.
its
Gadow's conclusions
Palceognnthce judged by this standard.
He
relation to the middle mesenteric
as to
remarks
:
the primitive nature of the
—"
It is
plain that,
so
far
as
PHrL0(5ENY OF THE PAL.llOGNATH^ AND NEOGNATH^.
249
degree of divergence of type in the alimentary canal goes, the Ratites deserve
their
accepted place at the bottom of the avian scale."
have been
I
So
Sfruthio.
enabled to
my
far as
examine the
material goes
it
intestines
of
all
the Palmgnathce, save
entirely confirms the observations of
Gadow,
and Beddard.
Mitcliell,
Casuarius,
Dromaus, and Apteryx
Droimms
In Casuarius and
resemble one another rather closely.
the duodenal loop is wide, in Apteryx narrow.
all
In
Casuarius, as Mr. Mitchell points out, the pancreatic and liepatic ducts open into
a
diverticulum of the di.stal loop.
In Dromaus and Aptenjx the duodenum is longer and
narrower, and
pancreatic and hepatic ducts ojien separately into the duodenum.
genera the small intestine is relatively sliort, and the convolutions therefore
In other
tlie
The rectum is short and straight; in Casnarins and 7>romr*'?fs its inner
thrown into a number of strongly marked irregular rugous folds.
In Casuarius and Dromaus the ca^ca are relatively short, of a uniform thickness
throughout, and have a narrow lumen.
In Apteryx the caeca are relatively much
are
simple.
lining
is
longer, exceeding
inches in length, whereas in the two former they do not exceed
Furtliermore, in Apteryx the terminal half becomes suddenly and markedly
5 inches.
dilated
each caecum thus presents a sausage-sliaped free end, borne on
;
a
narrow tube-
like stem.
There are no valvular appendages
lihea
is
in the caeca of tliese three forms.
intermediate between Casuarius on the one hand and Sfruf/uo on the other.
The duodenal loop
narrow and
is
resemble Casuarius, the posterior rectal
The resemblance
ducts open separately.
length of the rectum
and not a
The
series of
but
;
it
They
differs
to
of the small intestine
pancreatic and hepatic
Struthio consists in the relatively o-reat
therefrom in that
it is
only thrown into a single,
complex loops.
cd'ca are relatively of
(fnatJte.
The convolutions
portion Strutliio.
The
closed.
enormous length, exceeding those of any other Pahvo-
however, to vary in this respect very considerably.
a))pear,
In the
measurements given by Remoucliamps [85] the variations in li. mavroin R. americana between
rliyncha range between 1 foot 4.} inches and 2 feet 9 inches
In a female of this latter species dissected by me
2 feet 4 inches and 4 feet 8 inches.
series
of
;
they attained a length of 5 feet o inches, accompanying the small intestine throughout
the greater pari of
its
length.
The lumen was
sacculated to within a short distance
of the extreme end of the caecum by means of a series of semilunar valves alternatino-
one with another.
The
caecum by means of a
The
intestinal
therefore append
position of these valves
coils of Struthio I
outside of the
tlie
have not had the good fortune
Mr. Mitchell's description
of the intestine and
was indicated on
series of constrictions (fig. 7, p. 2-50).
the
three
[60].
to
He writes: — "The
great veins were according
to
ty])e,
examine.
first
the
I
two parts
yolk-sac
diverticulum being conspicuous in the circular loop opposite the end of the median
ME. W.
250
P.
PTCEAFT ON THE MOEPHOLOGT AND
Fig.
Caeca of
Bhea americana.
s.r.= small intestine.
c.=coeca.
Z.i.=large intestine, forming r.?.=rectal loop.
The
mesenteric vein.
distal
limb of the duodenum presented a short lateral diver-
ticulum, and the caeca are relatively longer
than in
Casuarius,
and, as
described frequently, were marked by the attachment of a spiral valve.
part of the intestine, that supplied by the posterior mesenteric vein,
an enormous
mesentery.
coil
Only
swung
the
at
in CJiauna,
circumference
has been
But the
is
rectal
expanded into
of a semicircular expansion of
and in the Eagles and Petrels, have
I
its
found the slightest
trace of a convergent resemblance to this feature, but in the latter the subsidiary rectal
loops, although supplied
According
These
caeca,
rectum by a
The
by the rectal
vessel, lie
above the
cseca."
Beddard the
spiral valve of the caeca contains about twenty turns.
he points out, are furthermore remarkable in that they open into the
to
common
orifice,
thus differing from those of
all
other birds.
Tinamous resemble those of Casuarius
Calodromas elegans,
however, affords a remarkable exception, differing as it does in this matter from all
cfeca
of the
;
known birds. These caeca have been described and figured by Beddard [6].
They are much wider than usual, and have the outer walls produced into numerous
other
prominent diverticula decreasing iu
size
from behind forwards.
—
PHYLOGENT OF THE PAL^OGNATH.E AND KEOGNATHtE.
The
following table from Beddard [7] will give at a glance the
the proportions of the alimentary canal
known
251
variations in
:
lih.i'u iiuicrorJii/ticJiii,
$
lUiea mnrrlcaiKt
(young)
Struthio camcliis,
!J
?
I,
„
?
,1
c^
d
CasuKfiiis inuq)/)!
iiilicnJitlus,
2
hi'iuani,
2
.,
casiicij'iiis
,,
picticulUs,
d
,,
hu-iinmcnhtiii.i
„
hennctt'i, c?
T)fOincn(f norti-JioUandli
Ajitcnj.i- ausiridix
oiveii.i,
„
(S
The
The
liver,
in
Neognatlue in that
all
the
it is
Palceognathce,
Liver.
save
that
of
the
is
larger tlian the
left.
A
Spigelian lobe
represented.
Both Casuarius and Dromceus possess a
in
with
divided into two lobes.
In Casuarius and Droiiueus the right lobe
is
agrees
Struthio,
common
In the former the gall opens,
gall-bladder.
with the pancreatic ducts, into a diverticulum on the distal loop of the
duodenum.
In Apteryx the
left
liver-lobe
is
larger than the right
;
the gall-bladder
is
occasion-
ally absent.
In Bhea
the right and
generally wanting.
left
liver-lobes
are equal in size.
The
gall-bladder
is
MR. W.
252
Stnithio
is
PTCEA^FT
P.
ON THE MORPHOLOaT AND
remarkable in that, according to Beddard, the right and
There
fused into a single heart-shaped lobe.
is
wanting
;
and there
is
is
a Spigelian lobe.
left
The
lobes are
gall-bladder
but a single bile-duct, which opens near the pylorus.
Mr. Beddard found the free edge of the
Bhynchotus mfescens pierced by
liver in
three small vessels of the vennl portal system
—
a fact whicli, as he points out, has a
very lizard-like appearance.
RESPIRATORY ORGANS.
a.
The Lungs (PL XLV.
The costo-pulmonary muscles
in all
on
;
in
fig. 9).
Casuarius are short, thick, and powerful, five pairs
they arise from the thoraco-sternal articulation of the ribs and extended fanwise
to the
Those of Rhea are
aponeurosis covering the lung.
relatively longer than in
In Apteryx, according to Huxley [41], they form " broad flat bands," which
Casuarius.
" take their origin from the vertebral ribs, at some distance below the attachment of
the pulmonary aponeurosis, and, proceeding obliquely dorsad and forwards, spread
out and are inserted into that part of the aponeurosis which covers the posterior
inferior facet.
In the relative size and distribution of the ecto-, ento-, and para-bronchia, and in the
size of the vestibule, the flightless members of the Palceognathce do not appear to
differ
much one from
another or from the Neogiutthd'.
In the lungs of a Casuarius unappendiculatus the bronchial rings extended backwards
as far as the 4th ento-bronchium.
The inner border
of the anterior and posterior
lung are deeply sculptured by a system of numerous parallel
inferior facets of the
grooves of open tubes, closely packed and running along the surface of the lung from
the mesial border outwards so as to converge towards, and open into, a spacious
communicating
chamber
(PI.
XLV.
fig. 9),
tlie
ventrally
with
the
1st
ento-bronchium
These grooves give the ventral face of the lung,
after
the
aponeurosis, the appearance of having been bored by Teredos, the groove
representing a heraisection.
tissue
and
further forwards and also ventrally into the anterior end of the
pre-bronchial ostium.
removal of
posteriorly
They
are crossed and recrossed by transverse strands of
and punctured by minute apertures.
They may be divided
into two groups according to
whether they are fed by a trunk
Thus, as will be seen in fig.
secondary grooves cephalad of the main trunk marked " ent. I." are fed from
from the
1st or the
ento-bronchium
III.
iJrd
ento-bronchium.
9,
all
the
this, whilst
supplies all the grooves caudad thereof.
These grooves are converted into tubes by the pulmonary aponeurosis.
b.
The
Air-s.\cs
In Casuarius the p-e-hrovchial sac
is
(PL
XLV.
paired,
fig. 8).
and extends forward on each side
of
—
PlirLOGENY OF
the oesophagus for a
thoracic
The
PAL.EOGINATH^ AND NEOGNATH^.
TilE
considerable
beyond
distance
the
of
level
the
2-33
cervico-
1st
rib.
suh-hronchial sac
divided into right and
is
loculi, enclosing a median
formed by the trachea. This
tunnel can be traced into a large pneumatic cavity in the sternum openin<^ on its
anterior border in the middle line between the coracoid grooves.
tunnel-like space between them, the roof of
The intermediate
scribed by
it
Beddard
would seem that
were three in number.
sacs
in
which
left
is
A
similar subdivision has been de-
Podargus and " one or two Accipitres."
In
tlie
present instance
due to a subdivision of the anterior intermediate.
The i^osUrior intermediate was large. Its roof was pressed inwards by the forward
this third sac is
extension of the posterior abdominal.
The
posterior abdominal
the right kidney
Avithout the sac
;
on the
and
left side
ostium from ento-bronchium
tlie
right side were markedly
:
—The
pre-bronchial and sub-bronchial by a
common
the anterior intermediate («) from ento-bronchium in.,
from the anterior of two canals arising one immediately behind the other from
the ventral floor of the
ecto-bronchia.
III. -IV.
lie
left.
sacs are fed as follows
(5)
side enclosed
the anterior lobe of the kidney was found to
posterior abdominal sacs of
smaller than those of the
and
That of the right
size.
to be enclosed in a separate sac.
The intermediate and
The
was of considerable
ecto-brouchial into
tlie
i.
;
meso-bronchium immediately below the apertures of the
The ento-lu-cnchial ostium opens into the anterior, the
posterior loculus.
of the two hinder ecto-bronchial canals.
—the odium
end of the meso-bronchium
The
The
posterior intermediate by the hinder
posterior
abdominal by the terminal
jiosterius.
The 2ud ento-bronchium does not open,
as described by
Huxley
in A'pteryx, into
the sub-bronchial ostium, but turns upwards and backwards, and terminates ciEcally
like the 4th.
The air-sacs in lihea differ in no essential respect from those of Casuanns.
The sub-bronchial sac is unpaired. In one Ehea americana which I dissected
anterior intermediate sac
was fed by two
ostia widely separated
a second example the ostium of the anterior intermediate sac of
septum dividing
The
from the
diff"ers
from
all
much
first
pointed out
to their smaller size, the posterior intermediate
In
all
inter se
l)y
Huxley, in
sac,
so that
tlie
it
projects,
abdominal
— PART
v.
like
a
cavity.
organs of the Pakeognathce are singularly uniform in character
and with the Neognathce.
VOL. XV.
that,
and abdominal sacs are enclosed
hernial sac, between the peritoneum and the parietes into
respiratory
and the
other birds a great increase in the size of
two has driven out the posterior abdominal
The
side
larger than those of the Cassowary.
other birds, as was
within the sub-pulmonary chamber.
tliese
tlie left
In
jiosterior intermediate W(>re l^otli wanting.
posterior abdominal sacs are
Aptcryx
owing
this sac
the
one from another.
No. 19.
December, 1900.
2 n
ME. W.
254
PTCKAFT ON THE MORPHOLOGY
p.
The
c.
The most valuable of the numerous
—from a systematic point
of view
—
AVindpipe.
on the windpipe of the Palceognathw
treatises
We
that by Forbes [20].
is
AjN'I)
are concerned here
only with the thoracic end of the trachea, and the degree of specialization which
it
has
reached in the direction of the formation of a " syrinx."
Struthio and Apteryx, up to the time of Mr. Beddard's paper [8], have been held
to represent the least specialized forms in this particular.
There
In Struthio there are no intrinsic muscles.
Forbes, " the
mucous membrane of the
interior
greatly thickened, forming a vocal
is
cord, in the region of the last three tracheal rings
The
membrana tympaniformis
" Internally," according to
a
is
but only a trace of a membrana semilunaris.
interna,
and
first
two bronchial semirings.
lateral tracheal muscles stop at the point of insertion of the sterno-tracheales!'
In Apteryx
membrana
there
is
semilunaris.
a broad
As
membrana
and a feeble
in Struthio there is
tympaniformis interna
no pessulus and,
it
was believed, no
intrinsic muscles.
Mr. Beddard's
howe\er, showed that
dissections,
these
were " especially plainly
In this species he found " a thin band of muscle
developed in Apteryx australis."
continuous above with the extrinsic muscle
and gradually thins out into fibrous
tissue,
;
this passes
down
which seems
the side of the trachea
away upon the
to die
last
tracheal ring, hardly reaching the bronchi.
Its state, in fact, is clearly one of degeneraand very mucli on a par with the state of the corresponding muscle in the
Hoatzin (Opisthocomus). In A. mantelli the same muscle was visible, but it did not
seem
to be anything like so well developed as it is in the species first referred to."
tion,
It appears to be entirely
There
is
wanting in A. haastii and A.
a feeble vocal cord,
and the
lateral
oiveni.
muscles stop at the insertion of the
sterno-tracheales.
Casuaritis, as
Forbes remarks, shows one interesting peculiarity, the lower end of
the trachea becoming slightly
dilated
tracheal rings are incomplete behind, the
filled
up by
this,
after death,
fibrous
and
elastic tissue
above the
Furthermore, the
bronchi.
median space
to
which
this gives rise
By
arranged transversely.
last
being
the contraction of
the ends of the rings are drawn inwards, so forming within the
tube a low longitudinal ridge, resembling the tracheal septum
of
certain
Petrels
and Penguins.
There
is
no pessulus, the membrana tympaniformis interna passing over from one
A
bronchus into the other.
line
between
remarks, there
cord
is
the
is
two
very slight thickening of this
bronchi suggests a memh-ana
membrane
in the middle
semilunaris, but,
nothing that can properly be called by this name.
as
Forbes
The outer
vocal
well developed.
Forbes describes, but so
far
I
am
not able to confirm
this,
"
On
the internal wall of
PHTLOGENY OF THE PAl-^EOGNATHiE AND NEOGNATHiE.
the bronchus
much
a second
slighter
and
prominent
less
255
concave
fold, slightly
somewhat obliquely backwards and downwards, and supported by
forwards, running
the anterior ends of the third and sixth bronchial semirings."
There are no
expand
The
intrinsic muscles.
meet
so as to
The
sterno-tracheales in Casuarius and in
middle line in
in the
Bromrfm
front.
tracheo-bronchial region of the trachea in the adult Bwrnceus has not yet been
In a young specimen dissected by F(n"bes there seemed
described.
differences
between
it
be no great
to
The memhrana semilunaris seemed
and Casnarius.
to
be better
developed.
The
trachea of Dromceics
peculiar, in that its front wall is pierced by a long,
is
median aperture, caused by the deficiency of certain of the tracheal
region.
The lining membrane of the tube projects from this opening
vertical, slit-like
rings in this
and forms
a large dilatable sac,
which can be
filled
with
air at
It
will.
is
probably
concerned with the production of the drumming sound which this bird occasionally
Init seems to have
It has been described in detail and figured by Murie [63]
makes.
;
been
first
discovered and described by Peter Fremery [21] in 1819.
In Apteryx,
it
will
be remembered, the syrinx
The
muscles are degenerate.
by Forbes [20].
The
intrinsic muscles
last four or five, or
completely to form a box.
is
The number
even
but feebly developed, and the intrinsic
of Bhea were
six,
first
correctly described
tracheal rings fuse
more or
less
of rings involved in this fusion can generally
be traced by the interannular sutures, which appear never to be completely obliterated.
There is a complete and cartilaginous pessulus and a memhrana semilunaris. Forbes
describes,
and
I
am
able to confirm,
"on
the middle part of the
formis interna a thickened fibro-cartilaginous area."
base resting on the pessulus, and
its
This
membraua tympani-
triangular in form,
is
its
apex terminating near the 4th bronchial semiring.
thickened tissue with certain fibrous peashaped pads found in this region in Sula hassana, first described by Garrod, and
Catarrhactes chrysocome, described and figured by Watson in the Challenger' Reports.
The broncho-trachealis (intrinsic syringeal) muscle in my dissections, as in Forbes's,
It
would be interesting
to
compare
this
'
arises
from the
and passes downwards to be inserted into the
bronchial semirings and the membrane between.
last ten tracheal rings,
anterior ends of the last four or five
The sterm-trachealis terminates
just pre-axiad of the origin of the intrinsic muscle.
In the Casuariidce, as Forbes has pointed out, the " terminal part of the trachea tends
to develop into
an expanded tympanum "
;
the tracheal rings are softer and
imperfect posteriorly, the junction of the trachea and bronchi
arrangement of the sterno-tracheales is peculiar.
In Sfruthio and Apteryx the tendency of the trachea
is
to
termination, and the distinction between this and the bronchi
less
is
become
marked, and the
become narrower
is
much
at its
sharper than in
Casuarius.
2n
2
ME. W.
256
P.
d.
The choanse
Avill
The Choanjs
of the Pakeognatha',
Neognathoe in their larger
as
PYCRAFT ON THE MORPHOLOGY AND
size.
In
XLV.
fig. 1).
interesting to remark, differ in form from the
it is
all
(PI.
the Neognathce the aperture
be seen in the figure, in the PakeognathiV
it
is
slit-like, whilst,
triangular or cordiform in
is
shape.
CIRCULATORY SYSTEM.
The Heart.
The
heart of the Pakeognathce does not differ materially in structure from that of
the Neognatho:;.
pointed out by
has been already ably described and
It
Owen
[67],
so that little remains for
Lankester [45, 46], Beddard
me
its
[5],
most important features
Rolleston [87], and others,
to say.
Fig. 8.
A.
P
^.se;.
fj.Z^.
7HO<l 6
/:c/u.
A.
it,
Dissection of the heart of Casuariiis unappendciihivs, to
show the right auriculo-ventricuhir valve and
the " moderator baud.'"
]<.
Dissection of the heart of Ulna amer'mtna, to
show the
right auriculo-ventricular
septal valve.
acc.t.
=
accessory tendon.
sep.v.
=
septal portion of right auriculo-ventriculnr valve.
s.v. =
r.au.v.
2).
rnorl.h.
—
semilunar valves.
right auriculo-ventricular valve.
= papillary muscle.
= moderator band.
band and the abnormal
PHYLOGENY OF
My
RoUestou.
concerns
point
only
It
seems
was always present,
I
the
not in
moderator band
all
though
I
described
Casuarins h)
in
this
the Palwognathce, at least in the Casuariidm.
have only succeeded in finding
salvadorii,
first
257
been taken for granted by subsequent writers that
to liave
if
PAL.^OGNATILE AND NEOGNATH.E.
TIIF.
it
once,
and in the heart of a Casuarius casuarius
have examined specimens of
C. unapi)endic%ilntus,
casuarius
C.
and Casuarius casuarius.
beccarii, C. casuarius violicoUis, C. casuarius sclateri,
In addition to the large papillary muscle binding the right auriculo-ventricular
valve to the anterior wall of the ventricle, 1 find two or
tendinese
"
" chordae
more very slender
extending from the outer surface of the auriculo-ventricular valve to the
anterior ventricular wall, recalling the similar structures described and figui-ed by
Lankester [45] in the heart of Ornithorhynclins.
In Bhea and Droma?us only one of
these threads was present.
I have, however, only had the opportunity of examining
one JJrovueus for this purpose.
The
septal half of the valve in Casuarius
relatively
somewhat
opportunity of examining,
border
and, in the
larger,
differs
curved forwards
is
forming by
means a
this
at
thick and fleshy.
is
adult
(uily
from Casuarius
its
heart
in that its free
insertion towards
sort of pocket-like flap
In Dromceus
which
it
is
have had an
I
ventral or hinder
the floor of the left auricle,
on the wall of the inter-ventricular
septum.
In Rhea this same arrangement occurs
(fig.
again.
8 b) this was remarkably well developed.
In one adult which I dissected
The
valve
was very
thin,
and the
forward pocket-shaped extension into the anterior end of the ventricle was strongly
developed and was continued yet further in the form of a narrow
attached only at
its
slip
of muscle
ends.
In Dromaics, in addition to one or two delicate tendinous threads from the outer
surface of the auriculo-ventricular valve to the anterior ventricular wall, there were
numerous
valve,
fleshy strands of muscle, interlacing
and from
it
one another, running parallel with thu
to the anterior wall of the ventricle.
Trabeculse carnese were well developed both in
Bhea and Dromceus.
Carotids.
All the Palceognathoi, save Apterijx, have normally two carotids.
Rhea, the
left
only persists.
In Apteryx there
is
only one carotid, the
Occasionally, in
left.
CLOACA, BURSA FABRICII, AND URINO-GENITAL SYSTEM.
a.
The
is
Cloaca.
terminal end of the alimentary canal in the PaUeognathce, as in the Neognatha\
divisible into a copro-, uro-,
and proctodceum,
as defined
by Dr. Gadow
[24].
In Casuarius, Rhea, and Apteryx the coprodisum passes g radually into the rectum,
but in Struthio and some Neognafhce the coprodceum is preceded by an additional
258
MR. W.
chamber, though
this
chambers therefore,
is
P.
PTCRAFT ON THE MOEPHOLO&T AND
but small and not very sharply defined.
as Dr.
Gadow
In
this succession of
has pointed out, Strnthio resembles some Saurians
and young Crocodiles.
In Strnthio, defaecation and micturition are separate
passing from the uro- to the coprodseum,
unlike that of
Its
all otlier birds, is
is
acts,
of considerable size
and remains permanently open.
function, acting as a physiological dorsal bladder,
bladder, communicating directly with the ureters,
The
and the urine, instead of
transferred to the bursa Fairicii, which,
is
is
unique.
A
true allantoic
wanting in the Aves.
cloacal aperture of the PakeognatJue, as in Neognathce,
is
round, not transverse
as in Reptilia.
Fi?.
9.
^'/
The terminal end
Casuarivs vioKcoUis, to show the cloaca and
of the alimentary canal of
into copro-, uro-, and iirootodaeum
..
and the form and position of the bm-sa
its divisions
Fabricii.
= buraa Fabricii.
= coprodjeum.
/)./.
c.d.
u.d.
=
p.d.
= proctodeum.
urodceum.
= clitoris.
ovid. = aperture of oviduct.
d.
II.
=
b.
aperture of ureter.
The Bursa.
The bwsa Fabricii of the Palceoynathce has been
[20] and Wenckebach [97].
In the Palceognathce
bursa in young birds
is
— save
—
only in Apteryx
of considerable size.
as
very carefully studied by Forbes
Forbes [20 a] has shown, the
Furthermore, he held that
it
differed
PHTLOGENY OF THE PAL^OdNATH^ AM) NEOGNATILE.
fundamentally from that of the Neogitatha, since in these
or
constricted
less
Struthiones
"
From
aperture
into
the
will be at
it
opens by a more
general cavity of the cloaca, whilst in the
the exact opposite was the case,
this
last it
259
the
opening into the bursa.
cloaca
once evident that the cloaca does 7wt open directly to
the outer surface, but indirectly tlirough the bursa by means of
its
large posterior
and
inferior aperture."
I tind
myself unable to confirm this point, and I think the accompanying figure will
The only difference which I can see between the Palwo- and Neognathce in
justify me.
this particular is that in the latter the bursa
means of
minute aperture,
a
in
the former
opens into the roof of the proctodseum by
opens by a very large one, so large as to
it
absorb nearly the whole roof of the proctodaeum.
When
place in
these parts are dissected fi-om spirit-specimens, in which hardening has taken
situ,
Forbes's description seems to be amply verified, as I can testify.
accompanying figure was made from
had been removed soon
after
The function
tlie
left for
is
yet
c.
That of
very
all
that
of
tliree
days.
The
unknown.
C()Pui>AToRv Organs.
the
grooved organ containing erectile tissue and resembling
Chelonia
in the floor of the proctodaeum.
and
Crocodi/ia.
The
ni.
can be retracted into a
It
protractor penis receives a slip from
the transversus analis, which descends from the pelvic bones.
j)enis arise
wall of one side was
possess a well-developed copulatory organ.
fStruthio is a solid
nearly
pouch
two or
true relations revealed.
of the bursa
The Palavgnatlue
specimen in which the hinder end of the gut
death, the cloacal aperture set widely open, and the
whole plunged into formol and
then removed and
a
The
from the pelvis and are attached
A
pair of retractores
to tlie ventral aspect of the
middle portion
of the organ.
In the remaining Faloognatluf the penis resembles that of the Anseres amongst
the Carinatu', being evertible and more or less markedly spirally twisted and grooved
dor sally.
The copulatory muscles
of the Palceognatha', Dr.
from those of the Neogiiatluc, inasmuch as the
Gadow
latter are derived
solely, whilst in the Pahtognatlia- tliey are partly differentiated
still
attached to the pelvis, and
[24] points
out, differ
from the sphincter ani
from muscles which are
are therefore skeleto-genital.
DEVELOPMENT.
The egg
of the Paheognatha', save in Apteryx and perhaps
tlic
that of the Neognatlue in that the pore-canals are branched.
Crypturi, differs from
In Apiteryx, as in the
Neognathie, they are unbranched.
In Struthio, according to Nassonov [64], and Apteryx, according
to
Parker [71], an
—
ME. W.
260
p.
PYCRAFT ON THE MORPHOLOGY AND
opercular fold grows backwards from the hyoid arch so as to cover the
branchial
clefts.
and described
to
it
This has been noticed in some Neognathw.
The
in Falco timmnculus.
have been made by Rathke [84]
first
and second
JSushkin [94] has figured
discovery of this operculum appears
first
in 1832.
PHYLOGENY OF THE PAL^OGNATH^.
upon the discussion of
I enter
this subject Avith very considerable trepidation
only on account of the difficulty of the problem, but also because
direct conflict with the
who speak
opinions of those
after
it
brings
deliberation
—not
me
into
and long
experience, and are, for this reason, and rightly, accounted authorities amongst us.
It
now
is
generally admitted that the Palceoynathce must be descendants of an
Avian proto-cariuate form capable of
flight.
Parker [71] has given a number of characters which lend the strongest support to
this view.
To many they will be convincing. The most important of these are
:
"The
"
"The
The
fact that the skeleton of the fore
The presence of an
alar
membrane
limb
that of a true wing."
is
or patagium."
presence of remiges and tectrices majores."
alar
membrane
is
According
a structure of great significance.
the remnant of a once extensive scale-covered patagium.
As
to Parker,
it is
these scales gradually
became transformed into feathers the size of the patagium decreased.
As to the mono- or polyphyletic origin of tlie group there is much divergence of
opinion.
Newton and Beddard consider the Pala^ognathce
Gadow, and Parker polyphyletic. AVith these last I
to
be monophyletic
;
Fiirbringer,
also agree, but not entirely so.
There can be no doubt but that the Palceognathw represent a lower grade than the
IfeognathcB.
It is
which
(fig.
Nevertheless the latter are closely related thereto.
probable that they have been derived as a diverging branch from that stock
gave
rise
the
to
10, p. 265) is intended to
The
inasmuch
it
It retains its
bears
upon the
The diagram
this clear.
the palate of the Cri/pturi to that of Bhea and
form of the pelvis of Crypturi
support to this view.
arch in the Palceognathce
structure of the pelvic
as
and jEpyornitlddce.
general resemblance in the
Dinoririt/iidce lend strong
The
Dinornitliidce,
make
very close resemblance of
Dinornithidce and the
and
Elieidce,
is
of considerable interest,
relation of this group to the Neognathce.
most primitive form in Archceopteryx, Binornis, ^j^^oniis, Hesperornis,
Ichfhyoriiis, A2)teryx\
and
in the Crypturi.
pubes are retained throughout
converted into a foramen by
Strufhio the ischiadic fissure
life.
all
these the ischiadic fissure and free
In Neognathce the ischiadic fissure becomes
synchondrosis
is
In
between the ilium
and ischium.
permanent, but the pubes fuse in the middle
In
line.
—
PHTLOGEXT (W THE PAL.EOGNATH.^ AND NEOGNATH.E.
In Shea the ischiadic
Ijelow tlie vertebral
seems
the ischia, furthermore, meet one another
Thus Struthio
tliein posteriorly.
form of the pelvic arch.
in the
Thus
to be a stumbling-block.
systematists think there can be
specialized
;
column, and the pubes fuse with
and Bhea are each unique
(•^^rHfA/o
closed
fissure is
261
little
Newton
Prof.
question of
writes [65]:
"Some
Struthiones being the most
tlie
and therefore probably the highest type of these Orders"
{'"''
Batitce").
Others, in spite of the great auKniut of specialization wliicli Struthio has undoubtedly
undergone, would regard
this
form as the most primitive, not only of living birds, but
of the Palmoipiathic living and extinct.
According
coracoid,
foot,
on account
Fiirbringer,
to
and many of
its
shoulder- and arm-muscles, the form of
"gewisse Eingeweide,
Vogeln,
number of
of the great
its
primaries,
its
pelvis, two-toed
entfernen sie weiter von alien anderen lebenden
etc.,
der Regcl unter einander divergiren.
als diese in
its
Zngleich bietet Struthio
neben verschiedenen mehr vorgeschritten Specialisirungen progressiver und retrograder
Natur eine Anzahl hochst primitiver Charaktere
dar, die ihre tiefer als
Katiten stellen und durch welche diese sehr alte
Vorzeiten des Vogelstammes thun
Fiirbringer,
as
remcjved far from the Dinornithnlce, as well as
is
from other Ratites, by the structure of
i'ar
die anderen
Blicke bis in die friihesten
lasst."
Beddard considers "that Struthio
however, go as
Form
its
for
palate,
which diverges much."
he continues further on
He
does not,
" There are really,
;
however, not a large series of characters in which they may be fairly said to be more
"
some other groups
primitive than
Fiirbringer's contention that Struthio
I find
is
This last statement
is
evoked in answer
to
we have
to
really very priuiitive.
myself in agreement with Beddard
:
more than
this, I believe that
look to Broitueus and not to Struthio for the most primitive of living birds.
This opinion
allied forms.
is
based mainly upon a study of the bones of the palate in this and the
These
from some closely
derived.
The
The
in
Broma'm seem
to represent the
allied form, the palates of Struthio,
palate of Apteri/x
must be considered by
ground type from which, or
Rhea, and Dinornis may be
itself.
palate of Struthio compared with that of Dromceus will be found by no means
so fundamentally different as seems to have been supposed.
contended that the palate of
form of that of
The
The
Strut/iio
the contrary,
it
is
a specialized and, in some respects, retrograde
JDroinceu.s.
palato-pterygoid connection
palatine and the forward
considerably
is
On
longer,
both
e.N
is
similar in both.
tension of the maxillo-palatine processes are both
actually
and
relatively,
in
than
Struthio
Dromceus.
in
Struthio lacks palatine processes to the premaxilla, and has a degenerate, one might
Like all degenerate structures, however, it gives some
almost say vestigial, vomer.
certain signs of a former perfection.
to Struthio meridional iii the
VOL. XV.
— FART
v.
No. 20.
Thus, in a skull which in
vomerine limbs extend backwards
December, 1900.
this
to
paper
is
attributed
within a very short
2o
262
ME. W.
.
PYCRAFT OX THE MOEPHOLOGY AND
P.
If this proximal extension were carried back sufficiently far
distance of the pterygoid.
to enable the
vomer
to join the pterygoid the
resemblance to the Dromeeine type would
be really striking.
If
we turn now
to the palate of
Bhea, and compare this with that of Drommis, we
The
the differences more decidedly marked.
shall find
peculiar and almost complex
which obtain between the palato-pterygo-vomerine articulations have already
been described in these pages (p. 206), and are sufficiently distinct to render further
relations
comparisons unnecessary.
But the palate of Hhea
So similar as to render
it
curiously similar to that of TJinornis and the Tinamous.
is
probable that these are related more closely one to another
than to Bromceiis or Strufhio.
The
affinity
though upon
though " this
of
is
removed
far
palate,
its
has already been hinted at by Beddard,
to the DinornithidcB
In discussing the position of Struthio, he remarks that
different grounds.
the structure of
remote
Bhea
from the Dinornitliidce,
as well as
which diverges much,
it is
from other Ratites, by
not clear that
Itliea is so
the existence of an apparent homologue of the maxillo-nasal bone
;
somewhat
point of
is
a
Emeus."
striking likeness to
Bhea and Binornis, was so impressed
with the likeness between them that he proposed to unite them in the same genus.
Parker [73], however, strangely enough, remarks " I know of no character in the
"
I say strangely
skull of Bhea by which it definitely approaches the Moas
drawn
attention
been
to the points
Parker's
enough advisedly, for 1 feel sure that had
Nathusius, again, in studying the egg-shells of
:
to
which attention
is
directed in this paper he
would have grasped
its
significance
and
written quite otherwise.
About the palate
we
of JEpyornis
as yet
resembles the Dinonu'thidce, inasmuch
know
nothing.
as, just as in
I
am
;
but since in
inclined to place
The
palate of the
its
it
pelvis
and other points
have already pointed out
palatine
having shifted from
connection more or
The
less
skull
it
it
resembles
closely resembles LinornithidcB,
it
near this group.
Tinamous
is
truly
Lromceognathoiis, but approaches the Neo-
gnathine type, tending towards schizognathism.
as 1
its
there are no out-
In some other points
standing supra-orbital processes to the lachrymal.
Sfruthio
In the roof of
this group,
(p.
208),
contact
intimate with
it
In
closely
its
pterygo-palatine articulation,
approaches the NeognathcB,
the
with the body of the pterygoid to form a
its tip.
palate of Apteryx in the peculiar forked pterygoid and the complexity of the
relations
between
it
other Palceognafhce,
Here, again,
is
and the palatine and vomer seems
or,
to differ
markedly from
all
the
indeed, from all living birds.
a conclusion diametrically
opposed to that of Parker.
Whether
or
not I failed to appreciate the points of resemblance Parker claims must remain to be
seen.
This discrepancy
is
certainly remarkable, and will
seem
to cast
doubt upon the
PHYLOGENT OF THE PAL.EOGNATHzE AND NEOGNATH.E.
validity of
my
conclusions rather than
liis,
who know
for those
development of 4.f.ry^. and the skull of the
mnornithid<B
monuments which mark an epoch in the history of
this
his
263
monographs on the
will a^n-ee that
they are
subject.
skull of the Linornithidce he wrote
Kiwis are certainly
for the
:
"
The n,arked
The real affinities underlyin<.
striking similaritv of the bones of
the
shown by the
palate in the two forms."
skull of CasHarius,
In his work on the
between the Moas and
niost part adaptive
these differences are, however,
The
differences
mny be remarked
it
here, differs in no essential respect from
from Dromceus point to specialization, as,
tor instance, the development of the
casque upon the mesethmoid
(p. 199, PI. XLIV.
ih^ioi
fig.
3)
The
lJro,n<,us.
In whatsoever
and the small quadrato-jugal
skull oiLromceus,
it
differs
it
fossa.
would seem, must be regarded
as the most generalized of
living birds, at least in so far as the
bones of the palate are concerned.
Strnthio somewhat nearly approaches Dromwus
in the arrangement of these bones
Mhea, the 'Iinamous, the Linornithidce, and
probably the ^pyornithidce agree more
closely one with another than with Dromxeus.
Ai>teryx- differs
We have
now
from
the direction of increased complexity of
these part.
Struthio and Lronicens witli Casuarius opposed
to Rhen, Linornithidce
all in
and Cryptur,, and possibly ^pyornis.
These last we may further subdivide by
means of the pelvis.
This will separate Rhecc from the remaining
forms; inasmuch
as
Bhea the pelvis is long and narrow, with the post-acetabular
ilia meeting
m
in the
middle
line, as in
Apteryx.
In the Binornithidw, ^:Epyorni.s, and the
Tinamous the
separated by the long transverse processes of the synsacrum.
It
may seem
that this character of the pelvis
reflection will show, I tliink, that
it
is
is
probably not
to the category of adaptive characters, since
post-acetabular
somewhat
a
so.
It
are
one, but
does not seem to belong
Rhea and Apteryx
mation of the pelvis agree, though their habits are very
ilia
artificial
in the general confoi--
and both bear some
of /y/r/^^/^/o— similar in so far as the great length
and transverse
width are concerned. In the Tinnmidw, Linornithidce, and
^pyornis the pelvis owes
its great breadth to
(1) elongated transverse processes, and (2) the broad dorsal
similarity to
different,
that
plane
of the post-acetabular ilium.
The
pelvis of Struthio bears an
undoubted resemblance to the Linornithine form,
Linornis or ^pyorni.% the transverse processes bear the
pos(acetabular ilia away from all contact with the neural spines of the
synsacrum.
Fiirbringer's view with regard to the Lronueidce is not exactly
in harmony with
tlie views adopted here, but if lends some support
thereto nevertheless.
inasmucli
as,
like
He
Lromceus and Casuarius "bilden zwei sehr uahe verwandte Familien
etwas holier
sowohl
als
die Struthionidce, aber tiefer als die Rheidce stehen
von den anderen Ratiten
als
von
den
Carinatcn
says
welche
und im Ubrigen
eine
2o
entfernte
2
— —
ME. W.
264
Stellung einnehmen,'
p.
PYCRAFT ON THE MORPHOLOGY AND
kommt mir am
and adds: "Ihre Vorgeschichte
meisten dunkel
"
vor
By
teudency of the foregoing will no doubt have revealed
this time the
intention
is
suggest that
to
Bromceus and Casuarius
regarded as the most primitive of the Palceognathce
most probably the nearest approach thereto,
;
— the
Dromceidce
Its
itself.
—are
to be
and that the Struthionidce are
spite of the great specialization
for, in
which they have undergone, they undoubtedly retain many archaic characters.
The
BinornitMdce,
Eheidce,
yEypormthidae, and
Cryj)t)iri
The
nearly related one to another than to the Lroma'idce.
aberrant of
all,
but have
affinities
are
all
probably moxe
A^iterygidcp are the
with the Linoridthida'.
This implies that the Palceognnthce are to be regarded as polyphyletic
triphyletic.
p.
An
most
attempt to express this relationship
is
—probably
diagram
given in the
on
265.
I propose to discuss the details of this " tree " in so far as concerns the Neocjnathce
elsewhere.
Of
the Paheognathce I need only say 1 recognize 7 Orders,
viz.
:
Casnarii.
Struthiones.
Rhese.
Crypturi.
Dinornithes.
^pyornithes.
Apteryges,
Their relation to the other members of the Class
Class
may be
expressed as follows
:
AVES.
Subclass
„
I.
II.
AnCIIiEOKNITHES.
Neornithes.
Grade A. Falofognathce.
„
It
final
B. Neognathce.
would have beeu helpful here
to discuss the further evolution of this
end in the various genera and species, but this
the present contribution owing to lack of material.
indication of the writer's views on this subject.
is
group
to its
unfortunately impossible in
The diagram
will
give a slight
PHYLOGENT OF THE PAL^.OGNATH^ AND NEOfiXATH^.
Caprimalgl
265
Acclpllres
Ciconis
J
Columb»
Pterocles
Strlges
Steganopodee^
fj
ff
^- Alcid»
Catharta
Pflittaci
Lunlcolai
GaUl
^pyomlthes
Phylogenetic diagram showing the probable Relationships of the PaloiO- and Neognatlm.
=
Palaeognathine stock.
'^^
=
Blank branches indicate eutaxic forms amongst a diastataxic group
eu-
and diastataxic forms occur
in the
^
Eutasic forms.
same genus or famOy,
;
=
Diastataxic forms.
blank spaces in a branch that both
ME. W.
266
P.
PYCEAFT ON THE MOEPHOLOGT AND
SUMMARY.
The group of
which
to
which we
birds
originally given by
name was
this
collectively designate the " Ratitae "
Merrem
A^jferyx, then unknown), on account of the fact that
to tlie
They were "raft-breasted,"
sternum.
keel-breasted
all
is
the same as that
in 1813 (with the addition of
agreed in the absence of a keel
in contradistinction to
the Carinatse, or
^.
Nitzsch, Huxley, Newton, Sclater, and Sharpe have successively adopted Merrem's
subdivision of the class Aves
— adding the
Saururae,
unknown
to
Merrem.
Other workers, both before and since Merrem, have accorded to the " Ratitae " no
Sometimes
greater prominence than that of an order or suborder of the class Aves.
rank of
to
the
even this was considered too great a distinction, and they were reduced
a Family, including such forms as Bustards, Plovers, &c.
The
" Ratitae " of
Merrem were apparently regarded by
that author, as well as by
Huxley, Newton, Sclater, Sharpe, and other systematists, as a monophyletic group.
Fiirbringer, Gadow, and Parker stand prominently forward as advocates for a polyphyletic origin.
Gadow's views are admirably
[26].
set forth in his contribution to
he regards the " Ratitae
Briefly,
"
Bronn's
'
Thier- Reich
'
as a morphologically monophyletic group,
standing in the same relation to the class Aves as the Anthropomorphae to the rest of
the
Mammalia.
With
within
the
Beddard
is
We
view we entirely agree.
this
[7], later,
was done long
This
group.
venture further, and include the Crypturi
since
by Garrod
has shown a strong inclination to follow
suit.
[30]
and
Stejneger.
Thus, he says:
— "It
unquestionably to the Strut/i W7ies that they showed the greatest number of important
likenesses, so
much
indeed, that their inclusion in one great group with
so,
would be by no means an unreasonable way of disposing of
The Dromaeognathous
Consent to
this,
but a makeshift
palate
is
sufficient
justification
them
tliem."
for
such
an association.
however, renders inoperative the old term " Ratitae," which at best
;
for
many of
is
the Cariuatae are Ratite, whilst the Tinamous have a
large keel to the sternum.
We
propose instead the terms Palceognathce (Ratitae
(Carinatae
Gadow
— Crypturi).
The adoption of
Crypturi) and Neognathw
these surmounts the difficulty indicated above.
regards the Struthiones as a group of primitive forms more nearly repre-
senting Proto-Carinatae than any other living birds.
name
+
" Ratitae
Similarly
"
Their retention under the
he regarded as convenient rather than an indication of close
we regard
the
PaUeog nathte
common
affinity.
of
as the unsuccessful descendants
proto-
carinate forms, but retaining probably but few of their characteristics unaltered.
would be too much
of any of the
'
It
to regard any of the existing Neognathoi as the direct descendants
existing
or
extinct
Blanchard in 1859 substituted
Talceognathw
for
known
to
us,
but
it
seems not
Merrem's the terms Tropido- and Homalosterni.
PHTLOGENY OF THE PAL.EOGXATH^
improbable that their source may be traced
NEOGNVTH.E.
AM:)
which gave
to that stock
267
rise to tlie
Rheo-
Dinornithine type of palate and pelvis.
Casnarius, which forms
nucleus of this research,
the
These two forms
generically distinct irom Droma'u.'f.
Strutldo
allied.
ever, these
here regarded
is
only
are, it is here contended, closely
perhaps derived from the same ancestral stock.
is
as
Hitherto, how-
two forms, Casuarius and Lromceus, have been regarded as more closely
Linomis and Apteryx than any other forms. Ajjferi/x we regard as quite
distinct.
The interrelationships of this group are indicated in the diagram (p. 265).
The skeleton of Brontceus (iter, procured at the last moment, at great trouble, by
allied
to
Mr. Rothschild, has proved of considerable interest
not
differ
in
one or two respects.
much, however, from that of the larger species: wherein
to approach the Cassowaries.
Skulls of adult Casuarius and Dromceus are badly needed
skeleton of EJiea darivitii.
tarso-metatarsus of
(tig.
darwi/ii
as
Z>.
also
irroratus.
is
the adult
temporarily reproduced in the embryo of R. ainericavn
four species of Ostrich indicated in this paper will be further confirmed
a point which time alone will sliow.
I
;
appears
interesting to note that the characteristic feathered
is
is
differs it
2 D, p. 155).
Whether the
is
Ii.
It
it
have seen no authenticated skeleton of
I
It does
hope
to
be able
to
supply
much
It
that
seems probable.
is
lacking in this paper by a further contri-
bution at no distant date in the sliape of an Appendix.
British
Museum
has acquired a hind-limb of Genyoniis.
account of the fact that digit
ii.
of great length and slenderness.
was becoming absorbed.
In this we
may
As Messrs.
Genyornh was undoubtedly a near
Dromceus.
Key
This
is
remarkable chiefly on
The proximal phalanx was
trace the early stage in the evolution
of a second two-toed Struthious bird.
ally of
Since this was written the
Stirling
and Zeitz have shown,
to the Osteology of the Pal^ognath^,
based on the Characters of the Adult Skeleton.
A. Skull.
The pterygoid
free anteriorly or resting
(Plates
XLII.-XLIV.)
ou the paired posterior ends of a large vomer
outer border with
tlie
;
couuected
mesial border of the palatine
with the palatine either by the opposition of its
op/josed glenoid surfaces.
or by squamous suture, or by complete fusion, but never bi/ means of
iu between
The vomer is always present, and jjaired, and is either fused with or tightly wedged
the maxillo-palatine processes
;
basipterygoid processes always present, of great
size, seated at
extreme proximal cud of the
the base of the parasphenoidal rostrum, and articulating with the
mesethmoid
processes of the pnmaxiUa unpaired; parasphenoidal rostrum and
pterygoid; nasal
continued forward beyond the level of the lachrymal.
A. Beak short or of
a.
Vomer deeply
medium
length, never probe-shaped,
cleft posteriorly
Into widely
expanded right and
running directly backwards beneath the pterygoids
;
left
limbs
palatine articulating
;;
MR. W.
268
MORPHOLOGY AND
Pi'CEAPT ON THE
P.
posteriorly by its mesial border with external border of the combined vomer
and pterygoid; vomer of great length, continued forwards beyond the
free
end of the paraspheuoidal rostrum
pterygoid
;
laminate
;
orbital
process of quadrate decurved and articulating with the pterygoid at
base
its
anterior palatine foramen very small, and lying near the tip of the
;
beak beyond the vomer; anterior narial and lachrymo-nasal
dried skull, confluent
fossje, in
mesethmoid with a truncated anterior border
;
nasal process of premaxilla rod-shaped
;
nasal with a vestigial external
process.
Vomer expanded
d.
palatiues,
into a broad flattened
plate
between the maxillo-
beak
and separated from the paraspheuoidal rostrum;
dorso-ventrally depressed
;
palatine fenestrated posteriorly, not anchy-
mesethmoid appearing on
but never inflated to form a
losed with pterygoid; horizontal plate of
the surface of the skull between nasals,
casque
;
lachrymals fusing with nasals and frontals
;
with long supra-
Bromxus.
orbital processes
V Vomer only very
.
slightly
laterally compressed
;
expanded between the maxillo-palatines
pterygoid and vomer;
;
beak
horizontal plate of mesethmoid, nasals, and
or less of frontals inflated to form a casque, sometimes of great
more
size
;
palatine unfeuestrated and anchylosed with the
quadrato-jugal fossa not extending forwards to level of antorbital
Casuarius,
pjfj^g
b.
Vomer
large, trough-shaped, closely investing the paraspheuoidal
rostrum
;
distal end of pterygoid and hinder end of vomer hidden (in ventral aspect
pterygoid rod-shaped.
of skull) by the palatine which underlies them
;
deeply cleft anteriorly, exposing the paraspheuoidal rostrum
between a pair of long spurs; anterior palatine foramen long and
Vomer
c'.
slit-like.
a".
Lachrymo-nasal fossa bounded anteriorly by a bony spike rising
upwards from the middle of the external border of maxillo-palatine
with large post-orbital processes
short, not extending
beyond
;
orbital process of quadrate very
level of articulation witli pterygoid
lachrymal with large orbital processes
;
mesethmoid
visible as a
Rhea.
lozcnge-sliaped plate between nasals
b".
Lachrymo-nasal fossa bounded anteriorly by
lating above with the nasal and
maxillo-palatine plate
;
a delicate rod articu-
below with the hinder border of the
no post-orbital processes
;
orbital process of
quadrate moderately long, with a free end projecting beyond articulachrymal without orbital processes ; nasal
lation with pterygoid
;
schizorhinal; inter-orbital
septum perforate; mesethmoid not
visible
Tinamida.
on surface of skull
d'.
Vomer truncated anteriorly, not extending so far forward as the end of
the paraspheuoidal rostrum; anterior palatine foramen small, closed
by anterior end of paraspheuoidal rostrum large post-orbital processes
lachrymals fused with frontals and not readily traceable; orbital
;
;
—
.
PHTLOriEXY OF THE PAL^OGNATH.E AND XEOGXATH^.
process of
pteiygoid
c.
iiiuuli'ate
long, articulatiuf;- «itli wliok'
inner l)ordcr witli
olfactory cavity of great size
;
Vomer deficient
269
Dinornithida.
not extending l)ackwards to reach the pterygoid,
anteriorly extending forwards, as in Dronuvas, to conceal the
parasphenoidal rostrum
the body of vomer deeply hollowed, fitting closely on to
postei'iorly,
;
the rostrum, and embraced on either side by the maxiIlo-|)alatine
processes ; maxillo-palatiue processes hastate
anterior jialatine processes of
;
great size;
pterygoid laminate;
palatine rod-sliaped, fu.sed proximally
with external border of pterygoid
prcmaxilla wanting a palatine process
lachrymals continued backwards to fuse with a chain of supra-orbitals
and with the frontals, so as to enclose a suin-a-orbital foramen lachrymal
;
;
;
with an osiculum lachrymo-]mluti»um; nostrils holorhinal
and
lachrymo-nasal
process bearing a
the middle of
its
fossa
confluent
well-defined
;
quadrate
projecting
facet
anterior narial
;
with
long
the
for
recurved
jiterygoid
in
internal ventral border
Strvthio.
B. Beak elongated, dccurved, probe-shaped; anterior narial fossa minute, eonfined to tip of beak; lachrymo-nasal fossa cnoruious, extending
forwards to
within a short distance of tip of beak
processes; lachrymal vestigial
backwards to optic foramen
;
skull rounded, without post-(n-bital
;
olfactory
chamber of great
vomer passing backwards
;
size,
extending
to fuse with the
pterygoid and palatine, and, with these, helping to form the floor of a
tubular recess opening inwards
Key
Aptenjx.
to the Species.
Struthio.
a.
Maxillo-palatine processes
not rising
dorsad, the
anterior
border of the
maxillo-palatine process at right angles to the long axis of the skull,
a'.
Palatines projecting distad beyond
process
anterior
border of
maxillo-palatine
maxillo-palatine processes without any external lateral fenes-
;
trated plate
b!.
s. aust rails.
Palatines not reaching as far forward as the level of the anterior border of
the maxillo-palatine process.
Vomer
a".
very short
;
parasphcnoidal rostrum thick
;
maxillo-palatine
processes with an external lateral fenestrated plate
Vomer
b".
very long
;
fenestrated plate
with
;
parasphcnoidal rostrum slender
deep ilepressions
meeting
in
S.molybdophaue.i.
maxillo-palatine processes without an external lateral
;
roof of the skull
mesiad of the supra-orbital ledges
the middle line proxiniad of
;
nasals
the nasal process of prc-
niaxilla
b.
S.meridioua/is'
Maxillo-palatine processes rising dorsad
anterior border of maxillo-palatine
;
processes sloping obliquely backwards
;
vomer broad, with
short, pointed,
posterior limbs; palatines projecting distatl beyond maxillo-palatine processes
'
S. camelus.
We
VOL. XV.
believe that Strutldo mas.iamis (^Noum.)
— FART
V.
No. 21.
is
identical with this bird.
Dtcemher, 19UU.
—
\V. K.
& W.
P. P.
2 p
ME. W.
270
p.
PTCEAFT
0^•
THE MOEPHOLOGT AND
Apteryx.
Vomer
a.
long,
equal
to
or longer than
the
width across the paroecipital
processes at widest, notched anteriorly, and with
the median cleft not
A.
extending forwards beyond the middle
b.
Vomer less than width across paroecipital
a'. Vomer notched anteriorly, the metlian
the paired [losterior limbs;
australis.
processes.
anterior portion nearly as long as
paroecipital processes well defined;
basi-
temporal plate bounded antero-laterally by a pair of ridges passing
forwards into parasphenoidal rostrum
V Vomer
.
A. haastU.
.
imperfect anteriorly, median portion
much
shorter than the paired
posterior limbs.
a"
.
b".
Size not less than 7 inches
A.
Size not less than 6^ inches
A. oweni.
a. mantelli.
Rhea.
A. Temporal fossa more or
less
sharply defined and extending backwards to
meet the lambdoidal ridge; lachrymal with elongated dorsal processes
arching backwards over orbit.
Posterior nares markedly wider anteriorly than posteriorly
a.
rine cleft not extending backwards
lachrymo-nasal fossa;
;
anterior vome-
to the level of the hinder angle of the
length of quadrato-jugal fossa less than width
across beak measured across the maxillary
pillars
bounding the lachi'ymoR. americana.
nasal fossa anteriorly
Posterior nares not wider anteriorly than posteriorly
b.
cleft
;
antei'ior
vomerine
extending backwards to the level of the hinder angle of the lachrymo-
nasal fossa; length of (juadrato-jugal fossa equal to width across beak at
the level of the maxillary processes bounding the lachrymo-nasal fossa.
E. niacrurhyncha
'.
B. Temporal fossa not sharply defined and not extending backwards to lambdoidal
ridge; lachrymals with short orbital processes; anterior vomerine cleft very
R- durwini.
short
Drom^us.
Length not exceeding 7 inches.
«.
D.novii-- hollandix.
a'.
Post-orbital processes short
b'.
Post-orbital processes long, extending
{^-^ in.)
Size not exceeding 5 inches
b.
downwards
to within a short distance
D. irroratus.
of quadrato-jugal bar
;
roof of skull with a very conspicuous cerebral
dome
'
The
D.
specific characters of this species
have been drawn from a single adult
ate?-''.
skull, kindly lent
me by
Mr. Beddard from the Zoological Society's Collection.
"
This skull
is
from a skeleton
closely resembles that of
in the Florence
Museum.
The bones
of the palate are missing.
D. novii-JioJhtndio; differiug therefrom chiefly on account
of its
The cranium
very small size and
the prominence of the cerebral dome, whilst in the peculiar form of the inferior limbs of the lachrymal
it
;
PHTLOGENT OF THE PAL.EOGNATH^ AND NEOGNATH^:.
271
B. Vertebr.e (uot including Cryptiiri) K
All the pre-synsacral vertehrre arc lieterocceloiis and free
high neural spines
iuterceiitra are wanting.
;
A. Hyperapopliyses of axis well developed
diapophysial
synsacral vertebne develop
all tlie
;
lamellae large
;
cervical vertebrae short
and conspicuous
one lumbo-sacral
least
at
;
cervical
;
vertebra; sacro-caudal (post-sacral) vertebne well ossified.
a.
Hyperapopliyses of atlas
moderate
neural
;
of axis rather
s[)ine
high
hyperapopliyses of axis well developed; cervical vertehrse rather short;
neural spines of cervical vertebne well developed and greatly excavated
postaxially
mence
at
no neural spines
;
laterally bifurcating
;
catapophyses com-
about the sixth vertel)ra, and do not form a ring
much produced
well developed, not
;
hypapophyses
diapophysiul
postaxiad;
extending towards postzygapophysis, large and conspicuous
himella
cervical
;
ribs
moderately long and styliform; thoracic and lumbar neural spines
not
much
axially
extended, and not subccpud in
vertebra? raised in part,
more or
less, aljove
from post-sacrals
sacral vertel)ne not plainly distinct
;
Total
a' .
[lost-sacnd
;
sacral
;
and post-
expanded and separating the post-acetabidar
sacral neural spines laterally
ilium
height
ventral margins of aeetabula;
caudal vcrtebrse eight or nine; pygostyle cylindrical.
number
of vertebne fifty-four
eighteen; vcrtebrse with ribs nine
;
vertebr<e seventeen or
cervical
post-sacral vertebrse nine or ten;
;
pre-axial articular surface of atlas moderately notched dorsaliy
pophysis of axis
;
hypa-
much elongated; hypapophyses developed from
the
eighteenth or nineteenth to the twenty-third vertebra; a conspicuous
fossa
on each side of the pre-axial part of the neural spines of dorsal
vertebrse
post-sacral
;
vertebne
raised
dorsad
decidedly
of ventral
margins of aeetabula
Total
b'.
number
Dromeens.
of vertebne from fifty-five to fifty-nine
fifteen or fourteen
;
;
cervical vertebraj
vertebrae with (free) ribs eleven or twelve
sacro-
;
caudal (post-sacral) vei'tebrre eight or nine; pre-axial articular surface
of
atlas
moderate,
may become
a foramen
hypapophysis
;
of axis
extremely long; hypapophyses developed from eighteenth to twentieth
vei'tebra; fosste be>ide pre-axial part of neural spines very small
caudal (post-sacral) vertebr;i raised scarcely at
b.
Hyperapopliyses of atlas large
;
spines of cervical vertebrae generally
;
sacro-
;
less well
.
Casuarius.
hyperapophysis
cervical vertebras short
more or
;
dorsad of aeetabula
neural spine of axis lofty
of axis moderate or extremely large
resembles D, irroratus.
all
In the conspieuous development of the cerebral
neural
;
developed
dome
;
cata-
it
differs
from both
existinjf
species.
I would remark here that the specitic characters attributed to D. irroniliis are bated upon an cxaminatiou
of a single
marked
'
unnamed
to justify
This
Key
is
my
skull in the British
Museum
Collection
;
but these appeared to
me
to
be suihciently
determination of this skull as of D. irroratus.
compiled, with modifications, from Mivart's memoir.
2p
2
;;
ME. W.
272
p.
may form
pophyses
PYCHAFT ON THE MORPHOLOGY AND
a ring
extending towards
lamella
(liapo])liysial
;
postzygapophysis, large and conspicuous or extremely so
•
moderate or small
ventral margin of acetabulum
;
(/.
Total
number
of vertebrte
three
vertebrse
cervical vertebrae fifteen.
;
from
;
forty-five to forty-seven
hyperapophyses very long
;
uo bypapophysis
;
cervical ribs
;
post-synsacral
pre-axial articular surface of atlas scarcely at all
notched dorsally
to atlas
;
sacro-caudal (post-sacral) slightly raised dorsad of
to axis
minute liypapophysis
a
;
high
lu-ural spine of axis very
;
hyperapophysis of axis extremely large
;
catapophyses commence at
;
the fifth vertebra, in the tenth and eleventh vertebrse they form a
hypapophyses developed from the twelfth cervical to the third
ring
;
vertebra;
thoracic
ribs
stvliform
cervical
conspicuous
developing
spicuous,
small
exceedingly extended axially
d'.
ilia
;
postaxially
and lumbar neural spines
thoracic
;
and con-
extending processes
and subcqual in height; syusacrum with
sacral vertebr;e plainly distinct
ventrad of
extremely large
lamella
diapophysial
from post-sacral
;
post-sacral bent
free caudal vertebrae seven or nine
;
much
pygostyle conical.
Apteryx.
Hyperapophyses of axis moderate neural spines of cervical vertebrae
neural spines of
well developed and deeply excavated postaxially
;
;
catapophyses
and seventh cervical vertebrae bifurcating laterally
commencing at the fourth vertebra; hypapophyses
commence
fifteenth (last cervical)
fourth,
fifth, sixth,
at
vertebra; diapophysial lamella
extending towards postzygapophysis, large and conspicuous
;
cervical
ribs probably moderately long and rather styliform ; thoracic and
lumbar neural spines not much axially extended, and probably not sub-
equal in height
syusacrum with post-sacrals not bent ventrad of
;
ilia.
B. Axis vertebra with a hypajjophysis, with or without well-developed liyperapophyses ; cervical vertebne greatly elongated ; neural spines of cervical
catapophyses commencing about the
vertebrie small or almost obsolete;
sixth vertebra; catapophysis never forming a ring; diapophysial lamella
extending towards posterior zygapophysis, small and inconspicuous ; a
conspicuous fossa on either side of pre-axial part of neural spines either of
or
cervical
distinct
e'.
Total
vertebrie
thoracic
;
syusacrum with
sacral
vertebrae
plainly
from post-sacral vertebrie.
number of
with ribs
small
;
;
small,
;
cervical vertebra seventeen
;
vertebrae
pre-axial articular surface of atlas with a large dorsal
hyperapophyses of atlas moderate
neural spine of axis moderate
but not obsolete
neural spines aborted
(first
;
ten; no lumbo-sacral vertebrae; synsacrum with eight post-
sacral vertebra
notch
vertebrae fifty-six
;
thoracico-dorsal)
hypapophyses but
little
;
;
hypapophysis of axis rather
neural spines of cervical vertebrae
no neural spines bifurcating laterally no
hypapophyses developed from the eighteenth
;
;
to the twenty-first
(first
produced pre-axially
;
thoracic)
vertebra;;
a conspicuous fossa on
each side of pre-axial part of neural spines of thoracic vertebrae, but not
of cervical
;
cervical ribs long
and styliform
;
thoracic and thoracico-
Dtnornithida:.
—
:
PHYLOOENY OF THE PAL.EOGNATH^
lumbar neural spines but
A\l) NEOGNATILE.
extended,
axially
little
:
273
but considerably
extended dorso-ventrally and not subequal in height
synsacrum with
;
post-sacrals not at all raised dorsad of acetabula, but rather depressed
ventrad
jwst-sacrals yerfvdhj
saeral plainly distinct from
ossified
no himbo-sacral vcrtebrte; pre-eaudal vertebras
pygostyle a laterally compressed plate
;
;
post-sacral vertebrae;
ten
....
;
Sinithio.
f. Total number of vertebne forty-nine; cervical vertebrie fourteen; preaxial articular dorsal surface of atlas with only a minute dorsal notch
;
no hyperapophysis
atlas;
to
liypapophysis
neural spine of axis not very high
loped
;
fourth
neural spines of cervical
to
vertebra; almost obsolete,
neural
;
sixteenth vertebrje bifurcate laterally
last
cervical
to
;
a conspicuous fossa
;
spines
of thirteenth to
hypapophyses narrow and
on each
side of pre-axial part
of neural spines from twelfth to the fifteenth inclusive
short and not styliform
;
those of
hypapophyses developed from
second dorsal vertebra;
pre-axially produced
not very large;
axis
hyperapophyses of axis well deve-
;
vertebne abort
seventh
of
free cervical ribs nine
;
;
cervical ribs
synsacrum with post-
sacral vertebrae raised quite dorsad of acetabula, also very imperfectly
ossified
vertebrae
and much reduced
;
post-saerals nine
;
;
sacral plainly distinct
free caudal vertebne five
from post-sueral
pygostyle small
;
and cylindrical
The
Rhea.
vertebral formula
is
:
Synsac.
~^
DromiTus
CV. 18
:
Cv. Th. 3
:
Th. 3 +
1
+2
:
Lb.
(i
:
Eb.
sc.
-J
:
Sc.
2
Cd. 9
-|- !t
= .J5.
18
t;
Syusac.
~
s
^
Casuariits
Cv. 15
:
Cv. Tb. 4
:
Th.5-|-l+3:
Lb. 5
:
Lb.sc.2:
8c.
2
Cd. 9-|-9
= 5.5.
18
9
Synsac.
Struthio
Cv. 17:
Cv. Th. 3:
Th. 5-1-2:
Lb. 8
:
.Sc.
Cd. S-flO
2:
18
Sj'usac.
Uhea
Cv.l4:
Cv.Th.3:
Th.3-t-2+l:
Lb. 3
:
Lb.
sc.
4
Sc.
2
:
Cd. 9-1-8
= 49.
17
6
Synsac.
Aijteryx
(-4. a. manttUi.)
Cv.l5:
Cv. Th. 1
:
Th.4-|-3-fl:
"
»
8
'
Lb. 4
:
Lb.
sc.
4
:
Sc.
2
:
Cd. 3-f 8=45.
"
'
11
'
;;
MR. W.
274
PTCEAET ON THE MORPHOLOGY AND
P.
C. Sternum
The
sternal plate
is
keelless
(fig. 4, p.
217) and Pectoral Arciii.
the coracoid anchyloses with the scapula, and lacks the acro-
;
coracoid, the coraco-seapular angle
is
very wide
;
fuvcula vestigial or absent.
A. Coracoid grooves not widely separated.
n.
Metasternum pointed, no
posterior lateral processes
pro-coracoid not
;
forming a distinct bar.
a'.
Coracoid grooves and bases of coracoids overlapping
process long, directed upwards and backwards
;
;
anterior lateral
a pair of short recurved
processes lying on the anterior sternal border between the anterior
lateral processes
as
b'.
long
with vestigial clavicles
;
;
sternal plate nearly as broad
Dromceus.
coracoid longer than broad
;
Coracoid grooves not overlapping
short or olxsolete
no
;
clavicles
;
anterior lateral processes of sternum
much
sternal plate
;
longer than broad;
Casuarius.
coracoid as broad as long
b.
Metasternum more or
less pointed,
pro-coracoid of great
size,
with large posterior lateral processes
forming a distinct bar of bone
sternum with a conspicuous ventral prominence
;
no
;
body of the
clavicles
;
articular
Struthio.
surfaces for ribs of great width
B. Coracoid grooves widely separated.
a.
Corpus
marked ventral prominence
sterni basin-shaped with a
lateral processes absent
posterior
;
anterior lateral processes conspicuous, each with
;
a large pneumatic aperture at the base
articular
;
crowded together immediately behind the anterior
surfaces
lateral
for ribs
processes;
coracoid grooves at the base of the anterior lateral processes and separated
one from another by the wide emarginate body of the anterior border of
the sternum
b.
Sternal plate
;
coracoids long and slender
flat,
pneumatic
a'.
;
pre-coracoid relatively small
pneumatic
slightly or not at all
lateral processes always present
;
;
.
Rhea.
metastcrnal and posterior
facets for ribs not separated by deep
pits.
Articular facets for ribs widely spaced, and lying entirely behind the
anterior lateral processes, and not separated by deep pneumatic pits
coracoid neaily as broad as long
b'.
;
pre-coracoid not conspicuous
.
.
.
Apteryx.
Facets for ribs crowded, few in number, and partly underlying the base
of the anterior lateral processes
C. Sternal plate
flat,
pneumatic
separated by deep pits
;
;
;
coracoid extremely reduced
metasternal clement wanting
sternal plate
;
.
.
.
facets for ribs
exceedingly narrow from
before
jEpyornis.
backwards
'
The sternum and
Dinornithidce.
pectoral arch of the Cri/iiftiri will be found described on pp.
221 & 263.
;
PHTLOGENY OF THE PAL.EOaXATH.^ AND NEOGNATH^.
Key
275
to tlie Species.
Struthio.
a.
Middle region of anterior border straight or
groove
eoracoid
arched
;
length of
distance between articulations of sternal
than
greater
slightly
ribs 1-5.
a'
.
Posterior lateral pi'occsses long and spatulatc, with an extensively ossified
S. niolybdophanes.
metasternal region between the jjosterior lateral processes
b' .
Posterior lateral processes moderately long
between posterior
b.
;
Middle region of anterior border of sterniim sinuous
posterior lateral
S. camelns.
;
posterior lateral pro-
no projecting metasternal region between the
length of eoracoid groove less than the
processes
not spatulate
cesses
region
ossified metasternal
lateral processes small
;
;
distance between the articulations of sternal ribs 1-5.
c'.
Width between the
free
end
of the anterior lateral processes
much
greater
B. itieridionalis.
than the width across their bases
d'.
Width
across free end of posterior lateral processes about equal to the
S. australis.
width across their bases
Rhea.
a.
Anterior border of sternum between the eoracoid grooves slightly excavated
width between right and left eoracoid grooves greater than length of
articular surface for eoracoid
b.
articular surfaces for sternal ribs i
;
.
.
;
articular surfaces for sternal
'
Length 6^
in.
riljs
!•
.
.
.
Coracoid grooves distinct and the sternal protuberance but
Length not exceeding 3^
rio-ht side
R. macrorhyncha.
DrOMjEUS.
-D.
slightly developed
b.
R. amencana.
Outer border of sternum between the eoracoid grooves deeply excavated
width between right and left eoracoid grooves equal to or less than length of
articular surface for eoracoid
a.
.
in.
onlv represented
;
;
eoracoid grooves obsolete
;
the ventral
lip
novcB-hollmdia.
of
sternal protuberance conspicuous
D.
uter.
Casuarius.
ridges lying immediately behind a line
A. With a pair of low, median, keel-like
coracoid grooves, without
drawn across the sternum immediately behind the
C. wiappendiculatus.
grooves
median pneumatic cavity between the coracoid
the
between
cavity
pneumatic
large
with
ridges
B. Without paired median
;
eoracoid grooves.
a.
marked when seen from
Coracoid grooves deep, the lower lip strongly
backwards into a low
above; pneumatic cavity very large, extending
the sternum between
of
line
middle
swollen protuberance lying in the
the coracoid grooves.
ME. W.
276
p.
a'.
Width between inner
b'.
Width between inner
PTCEAFT ON THE MOEPllOLOGT AND
angles of articulation for last sternal rib less tban
width across metasternum at widest
C. casuarius.
angles of articiilation for last sternal rib greater
than width across metastcrunm at widest
C. c. beccarii.
lip not strongly marked when seen from
mouth of the pneumatic aperture produced forward into prominent
Coracoid grooves shallow, lower
b.
above
;
lips
anterior lateral process short and recurved.
;
Metasternal region
very long and narrow.
c'.
Width
d'
Width
across
anterior lateral processes
of sternum
width of
twice
C
metasternum
.
across
metasternum
C. picticollis,
D. Pelvic Akch K
Synsacral neural spines of great height
and
(PI.
XLIV.
sacral vertebraj,
;
fig. 5.)
and sometimes the
and second caudals with conspicuous pleurosteal elements
first
bennetti.
across anterior lateral process of sternum less than twice width
meeting in the mid-dorsal
spines never projecting
line
;
level
permanently open or closed only by the
of the post-acetabular
fissure of the
ilia
;
lumbo-sacral,
pre-acetabular
above the neural spines of the synsacrum
beyond the
last
;
ilia
always
the synsacral neural
ilio-ischiadic fissure either
extreme ends of the ilium and ischium.
A. Pubes remaining permanently free caudad.
With
a.
the post-acetabular
dorsal line; a
more or
Post-acetabular
a'.
spines
;
less
more or
less
widely separated in the mid-
well-developed supra-trochanteric process.
separated by the intervention of the synsacral neural
ilia
ischia
ilia
bowed inwards towards the middle
posteriorly with the ilium
;
and fused
line,
perforation of inner wall of acetabulum
constricted by ossification.
longer
a". Post-acetabular
than pre-acetabular
ilium,
and
tapering
gently
b"
.
DronKeus.
Post-acetabular
shorter
than
pre-acetabular
ilium,
and
arched
dorsally
Casuarius.
Post-acetabiilar ilia widely separated
b' .
by means of transverse processes,
leaving neural spines in middle of a deep fossa roofed in late in
a sheet of bone
post-ilia
c".
in,
flat
d'' .
;
Median
;
ischia widely separated one
life by
from another and from
post-acetabular with a very wide dorsal plane.
fossa dividing post-acetabular ilia only imperfectly roofed
leaving a more or less extensive cleft on either side of a Ijroad,
neural plate
Median
Dinornithidoe.
fossa dividing post-acetabular
but perforated by
a
ilia
completely roofed by bone,
double row of foramina, one on either side of
the median line
'
The
here.
pelvic arch of the Crypturi will be found described on
^pyornis.
]>.
2(i3.
There was no need to include
it
—
phyl()(;exy of the
b.
pal^ognath^ and neognath.e.
277
Post-acetabular
ilia closely approximated, the neural spines at most
only
barely visible between the closely approximated iunomiuates
no supratrochanteric process iseliia widely separated from post-acetabular ilium;
;
;
obturator foramen and fissure confluent
synsacrum welded into a large
laterally
;
post-sacral vertebra of the
compressed mass, bent down so
as to bring the neural crest to the level of the ventral border of the post-
''"i"
Apteryx.
B. Pubes auchyloscd caudad with one another or with ischium.
c. Post-acetabular ilia widely separated from iscliia, and from
one another
in the mid-dorsal line ; pubes fused caudad («) with the ischia and
(6)
with one another in the mid-ventral line
ing below the
d.
level of the ventral
Post-acetabular
ilia
mid-dorsal line;
;
the synsacral vertcbrce appear-
border of the ilium
Struthio.
anchylosed with ischia, and with one another in the
pubes anchylosed with ischia caudad; ischia clusclv
united in the middle line one wdth another throughout almost entire
the post-sacral vertebra; of the synsacrum atrophied
;
length (uuique)
.
Keij to
tlie
Rhea.
Species,
Rhea.
a.
Pre-ilium not projecting forward beyond the last thoracic vertebra and
which forms the
b.
limit of the
R, americana.
Pre-ilium projecting forward conspicuously beyond the
^^^
rib,
synsacrum cephalad
last thoracic
vertebra
^^^
R. macrorhyncha.
Apteryx.
A. Width across anti-troehanter equal
to or less than the width of pre-acetabular
ilium at widest.
a.
Pre-acetabular ilium of great width, with a strongly arched dorsal border,
and large
laterally
acetabular
ilia at
expanded antero-vcntral angles
widest part
= to
length of post-aeetabular ilium
widening at
free
its
end
;
;
width across pre-
length of ischium, and far exceeding
post-acetabular ilium with a sudden
breadth of pre-ilium at widest
;
= breadth
across
auti-trochanter
A.australis.
B. Width across anti-troehanter always greater than width of pre-acetabular
ilium at widest
widest
b.
less
;
post-acetabular tapering caudad
Pectineal process large
a'.
;
breadth of pre-ilium at
than breadth of anti-troehanter.
;
five
lumbar
vertebrse.
Post-aeetabular ilium with a concave ventral border, and a slight notch
immediately behind the anti-troehanter
b'.
Post-acetabular ilium with straight or convex ventral border
behind the auti-troehauter
c.
Pectoral process small
VOL. XV.
— PARTY.
;
No. 22.
four lumbar vertebrse
Decemher^Vd^^.
^. Uaasii.
;
no notch
^. oweni.
A.
a. inantdli.
2 q
;
ME. W.
278
PTCEAFT ON THE MOEPHOLOGT AND
p.
Drom.eus.
a.
Length 17
in.
mncli arched
pectineal process well developed; dorsal border of pre-ilium
;
;
obturator fissure nearly closed
;
piilio-ischiadic fissure nearly
D. novcE-hollandia.
closed
b.
Length 9i
arched
in.
;
process obsolete
pectineal
obturator fissure wide
;
dorsal border of pre-ilium not
;
pubo-ischiadic fissure very wide
;
.
.
.
.
D.
ater,
E. Pectoral Liaib.
A. Extended forearm and hand longer than the humerus
that of the typical Neognathte
chink between them
distal
;
B. Extended forearm and hand
Humerus with
a.
radius and
;
end of
much
IL
IMe.
ulnars resembling
;
ulna long, with but a narrow
Rhea.
flabelliform
shorter than the humerus.
a shallow, but blind subtrochauteric fossa; a very strongly
developed liuea aspera for the triceps, forming a conspicuous triangular
ridge
;
end of Me.
II.
not markedly expanded
typical carinate form,
pectoral crest
them
;
;
;
ulnare a simple nodule
;
manus
distal
;
as in Rhea, resembling
though too small for purposes of
Humerus much bowed, without
b.
bowed
radius and ulna short and
flight
....
radius and ulna with but a very narrow chink between
radiale and
a vestigial
ulnare
;
a vestigial pollex
an
bearing
unequal phalanx, with the Me. III. incomplete and free distally
.
.
C. Extended forearm and hand about equal to the length of the humerus
subtrochanteric fossa
a.
Subtrochanteric fossa large
;
;
Subtrochanteric fossa
Apteryx.
and ulnare anchylosed with metacarpals.
radiale
;
.
a
ulnare indistinguishable
radiale anchylosed
;
Casuarius.
with radius and carpo-metacarpus, but distinguishable
b.
Struthio.
of
subtrochanteric fossa or indication
very small
;
no carpal elements distinguishable
Dromaus.
neither pollex nor third digit represented
F. Pelvic Limb.
When
the bones are pneumatic only the femur
the extensor bridge
a.
is
wanting
Toes three in number
a'.
;
;
the hallux,
when
is
conspicuously so
present,
;
save in the Dinornithidee
tarso-metatarsus far exceeding femur in length,
Ectocuemial crest large, rivalling the procnemial in
a deep lateral pit
;
never large.
is
distal
size
;
euto-condyle with
extremity of shaft above entocoudyle (lateral
surface) with a deep groove,
fl".
Procnemial crest
extending
down
the
shaft
;
femur with a large
DromcBUS.
pneumatic aperture
b".
Procnemial crest not extending down the shaft
ungual phalanx of inner toe
b'.
gi-eatly
;
femur nou-pneuniatic;
Casuarius.
elongated
Ectocuemial crest small, a mere tubercle of the procnemial; no entocondylar pit or groove on the lateral surface of the shaft
conspicuous extensor groove
;
;
a moderately
popliteal fossa of the femur very deep
lingual phalanges laterally compressed
;
Rhea.
—
PHYLOGENY OF THE PAL.«OGNATH.E AND NEOGNATH.E.
Toes two iu number
b.
tarso-metatarsus far exceeding the femur in length.
;
Ectocnemial crest small, a mere tubercle on proenemial
c'.
to level of proximal tarsal
condylar
mass
;
fibula
extensor groove obsolete
;
continued backwards into a groove
pit
279
;
extending
a deep ento-
;
the lateral ventral
border of ento- and ecto-condyles continued backward and upward to
project spur-wise from the shaft
Toes four
c.
d'.
Struthio.
tarso-metatarsus short.
;
Extensor bridge wanting.
Femur long and
c".
slender; tibio-tarsus with a deep intercondylar gorge;
entocoudyle very large, projecting conspicuously in front of shaft in
lateral view
d".
Femur very
none of the phalanges conspicuously shortened
;
short
and thick, with enormous great trochanter;
clearly distinguishable
Extensor bridge
jjresent;
:
Ph.
D.
2.
I.
femur short
;
.
Apteryx.
.
tibio-
and ecto-condyles not being
tarsus without intercondylar gorge, ento-
d.
.
short
;
Ph. 2,
3,
1-.
D. IV. short
JEpyornis.
.
ento-condyle of tibio-tarsus large,
resembling that of Apteri/x
List of
1.
Ali.\, p.
H.
E.— Syrinx
Dinormthida>.
Works referred
to or consulted.
Bull. Soc. Philomat. 1874, p. 38.
of Rhea.
— On the Skeleton of the Apteryx. Joixrn. Linn. Soc, Zool. 1873,
Andrews, C. W. — Note on a complete Skeleton of /Ejjyornis from Madagascar.
2. Allis, T.
3.
xi.
vol. iv.
4.
Andrews,
p.
1897,
W.
C.
p.
p.
523.
Geol.
Mag.
Ibis,
1896,
241.
—On
the Skull, Sternum, and
Shoulder-girdle of jEpyornh.
376.
5.
Bedd.^rd, F. E.
6.
Beddard,
7.
Beddard,
8.
Beddard,
9.
Beddard,
F. E.
— On the Heart of Apteryx.
— On the Alimentary Canal
Proc. Zool. Soc. 1885, p. 188.
of the Martineta
Tinamou.
Ibis,
1890, vol.
iil.
pp. 61-66.
F. E.
vol. vi.
—The Structure and Classification of Birds.
— Notes on the Anatomy of the Genus
E.
F.
1898.
Apteryx.
Novitates
Zoologicse,
1899, pp. 38(5-402.
F. E.
and Mitchell, P. C.
— On the Structure of
the Heart of the Alligator.
Proc.
Zool. Soc. 1895, pp. 343-349.
— Abbildung.
—De Rhea americana.
10.
Blumexbach,
11.
BoECKiNG, A.
12.
BuRCKHARDT, R.
13.
Cunningham, R. O.
naturliistor. vol.
J. F.
Veher yUpyomis.
viii.
Diss. Inaug.
Bonn, 1863.
Palseontol. Abhandl. ueue Folge, Bd.
ii.
Heft 2 (Jena,
1893).
darwini.
—Notes
on some Points
Proc. Zool. Soc. 1871,
p.
in
the Osteology of
Rhea americana and Rhea
105.
2q2
280
14.
W.
:ME.
CuviEK, Baron G.
W.
15.
Dallas,
16.
DucHAMP, G.
PYCKAFT on the MOEPHOLOGy AND
— Leyoiis d'Anat.
— On
S.
p.
Comp.
1799-1800 (Ann.
t. ii.
—
viii.), p.
69.
Proc. Zool. Soc. 1865,
the Feathers of Dinornis robustus.
p.
265.
17.
Ann. Sci. Nat. xvii. 1873.
Observatious sur I'Anatoraie du Dromieus.
Edwards, A. Milne- and Oustalet, E. Notice sur quelques Especes d'Oiseaux du Museum
Ceuten. Mus. d'Hist. Nat. Paris, 1893 (I'fimeu on Emeu noir,
d'Histoire Naturelle.
18.
Evans, F. P.
19.
Flower,
p.
See also Milne-Edwards.
246).
p.
—
— Note
on the Carotids of Rhea americana.
Ann. & Mag. Nat. Hist.
xi.
1884,
458.
W. H.
Sir
— On the Skeleton of the Australian Cassowary, Casuarius
Proc.
australls.
Zool. Soc. 1871, p. 32.
20.
Forbes,
W.
21.
W.
Fremery,
the Conformation of the Thoracic
End
of the Trachea in the Ratite
Proc. Zool. Soc. 1881, p. 778.
Birds.
20A. Forbes,
—On
A.
A.
p.
—On the Bursa
J.
— Specimen
Proc. Zool. Soc. 1877, pp. 304-318.
Fabricii in Birds.
Zoologicum
sistens observationes,
pnesertim Osteologicas de
Casuario tiovie-Jiollandue.
M.
22. FiJKBRiNGEK,
23.
— Zur
Gadow, H.
vergleich. Anat. der
— Remarks
Gadow, H.
Amsterdam, 1888.
Muskulatur des Beckens und hinteren Gliedmasse der
Jena, 1880.
Ratitcn.
24.
— TJntersuch. znr Morphol. und System der Vogel.
on the Cloaca and on the Copulatory Organs of the Amniota.
Phil.
Trans. Roy. Soc. 1887, pp. 5-37.
25.
Gadow, H.
— On the Anatomical
1885,
Dift'erences in the
Three Species
(jf
Rhea.
28.
Gadow, H.— Bronn's Thiei-Reich. Bd. vi. Vogel, 1891, Anatom. Theil.
Gadow, H.— Ibid. Syst. Theil, 1893.
Garrod, a. H., and Darwin, F. Note on an Ostrich lately living in the
29.
Garrod, A. H.
— On the Carotid Arteries
30.
Garrod, A. H.
— On certain
31.
Garrod, A.
32.
Geofkroy
26.
27.
Proc. Zool. Soc.
308.
p.
—
Society's Collection.
Proc. Zool. Soc. 1872, pp. 356-362.
tion
Part
:
of Birds.
Proc. Zool. Soc. 1873, p. 457.
Muscles of the Thigh of Birds and on their Value
in Classifica-
Proc. Zool. Soc. 1873, pp. 626-644.
I.
H.—Ditto
:
Part
St. Hilaire, E.
II.
Op.cit. 1874, pp. 111-123.
— Compos, des Appareils genitaux, urinaires
points de rencontre dans I'Autruehe et dans le Casoar.
Mem.
et
d.
intestinaux,
Hist. Nat.
h.
leurs
tome
ix.
1822, p. 438.
33.
Gruber, W.
34.
Haughton,
1855,
p.
Acad.
35.
— Ueber das
Haughton,
S.
Thranenbein der Straussartigen Vogel.
Bull. Ac. Sci. St. Petersb.
161.
—On
vol. ix.
S.
the Muscular
Mechanism
of the
Leg of
the Ostrich.
Proc. Roy. Irish
1866.
— On
tlie
Muscular Anatomy of the Emu, Droiiueus iwvce-hoUundi(E.
Ibid.
1868, pp. 187-497.
36.
37.
—
Haughton, S. On Muscular Anatomy of the Rhea. Ibid. p. 497.
Hildebrandt, G. F. Bemerkung liber den Embryo des Strausses im
—
Naturkunde, Bd.
W.
38.
HuTTON,
39.
Huxley, T.
40.
Huxley, T. H.
F.
xi.
— The Moas of New Zealand.
H.— On
Eie.
1806, pp. 97-126.
Trans. N. Z. Instit. vol. xxiv.
the Classification of Birds.
Proc. Zool. Soc. 1867.
—The Anatomy of Vertebrated Animals.
1871.
Voigt's Magaz. d.
PlIYLOGEXY OF THE PAL.EOGXATH.E AND NEOGXATILE.
— On
41. Huxley, T. H.
42.
Knox, R.
Respiratory Organs of Jji/cn/x.
t!ie
— Observations
2f<l
Proc. Zool. Soc. 1882,
p.
560.
New
on the Anatomical Structure of the Cassowary of
Holland.
Edinl). Philos. Journ. vol. x. 1824, p. 132.
43. Knox, R.
of
— Additional
New
Holland.
44. Lankester, E. R.
Lankester, E. R.
Edinb. Philos. Journ. vol.
— On
Zool. Soc. 1883,
45.
Observations on the Structure of the Trachea iu the Cassowary
1823-24,
x.
137.
p.
the Right Cardiac Valve of Echidna and of Ornithorlujnchus.
8-14-,
j)}).
Emeu
Proc.
jils. iii., iv.
— Ou the Heart described by
Prof.
Owen
in
1841 as that of Apteryx.
Proc.
Zool. Soc. 1885, pp. 239, 240.
46.
R.— On
Lankester, E.
Owen
Richard
the Right Cardiac Valve of the Specimens of Apteryx dissected by Sir
Proc. Zool. Soc. 1885, pp. 4.77-482.
in 1841.
50.
— On the Avian Sternum. Proc. Zool. Soc. 1885, pp. 684-716,
Lydekker, R. — Cat. Foss. Birds, 1891.
Macalister, a. — On the Anatomy of the Ostrich. Proc. Roy. Irish Acad. 1867.
the New Holland Ostrich.
Macartney, — An Account of the Organ of Voice
51.
Malm,
52.
Marshall, W.
53.
Mayer, A. F. J. C. Analecten fiir vergleich. Anatoni. 1839.
Meckel, J. F. System der vergleich. Auatoni., Abtheil. ii. 1825,
Meckel, J. F. Beitriige zur Auatoraie des Indischen Casuars.
pis. xlii.-xlv.
47. Lindsay, B.
48.
49.
in
J.
Brit.
Assoc. Re]i. Bristol, 1836.
W.— Om
A.
1875,
54.
55.
CEfv. K. Vet.-Ak. Fr.rh. 1880, no. 7, p. 33.
Lnftror-sackeu.
— Beobachtungen
iibcr das
Verhiiltniss der Fcdern.
Gart. vol. xvi.
Zool.
121.
p.
—
—
p.
—
1830,
56. Meijere,
p.
200; 1832,
H.
J.
i)E.
p.
202.
Archiv
f.
Anat.
u.
Phys.
Bd.
4,
1895,
273.
— Febcr
Morphol. Jahrb.
Federu der Vogel.
die
xxiii.
pp. 562-591.
— Vcrgleichende
57.
Menzbier,
58.
Milne-Edwards and Grandidier.
59.
MiLNE-E,DWARns and Grandidier.
]\[.
Reiidus Acad. Sci.
t.
Osteol. des Penguine.
cxviii.
recents de Madagascar.
— Observations sur
62.
63.
64.
Imp. Nat. jNIoscou, 1887.
Bull.
— Sur
les
Ossenients d'Oiscaux provenant des Terrains
Mus. Hist. Nat.
pt.
1895,
p. 9.
See also Edwards.
C— On the Intestinal Tract of Birds.
—Ou so-called Quintocubitalism. Journ. Linn. Soc, Zool.
Trans. Zool. Soc.
MivART, St. G. —^On the Axial Skeleton of the Strut
405.
MuRiE, — On the Tracheal Pouch of the Emu. Proc. Zool. Soc. 1867,
[On the history of
razvitiya Afrlkanskagho Strausa.
Nasonov, N. V. — K
Mitchell,
Comptes
/Epyoniis de Madagascar.
1894, p. 122.
Proc. Zool. Soc. 1896, pp. 136-159.
60. Mitchell, P.
61.
Bull. Soc.
les
vol. xxvii.
P. C.
x. p. 1.
hionida'.
p.
J.
istoriT
lo])meut of the African Ostrich.]
Warsaw
University.]
3 Pt.
the deve-
[Published by the Zoological Section of the
1894-96.
— Dictionary of Birds (Introduction), 1896, 109.
1838.
Owen, Sir R. — Memoir on Anatomy of the Southern Apteryx. Trans. Zool. Soc.
145.
Zool.
Soc.
Trans.
R. — On Dinumis.
67. Owen,
68. Owen, Sir R. — Art. " Aves," Cyclopedia Anat. & Physiol. 1835-36.
Rep. Pur Seal Investigations, 1896-7, part
69. Palmer, W. — Avifauna of the Pribilof Islands.
1827.
Abtheil.
Osteologie,
—
Vergleich.
E.
Pander, C. H. and d'Alton,
Trans.
— Observations on the Anatomy and Development of Apteryx.
Parker, T.
65.
Newton, A.
p.
vol.
66.
ii.
vii. p.
-Si;-
3.
ii.
70.
71.
i.
Pliil.
J.
Roy. Soc. 1891,
Lief.
vol.
182.
ME, W.
282
72.
Parker, T.
J.
P.
—Additional
PTCEAFT ON THE MOEPilOLOGY AND
Observations on the Development of Apterijx.
Phil. Trans.
Roy.
Soc. 1892, vol. 183.
— On the Cranial
Osteology, Classification, and Phylogeny of the Dinornitfiidx.
73.
Parker, T.
74.
75.
—Text-Book of Zoology, vol
Parker, W. K. — On the Osteology of
76.
Parker, W. K.
J.
Trans. Zool. Soc. vol.
Parker, T.
1895.
tlie
Soc. vol.
77.
xiii.
J.
1897.
Gallinaceous Birds and Tinamous.
Trans. Zool.
1862.
V.
— On the Shoulder-Girdle and Sternum.
K. — On the Structure and Development
W.
Parker,
ii.
Ray
Soc. Publications, 1868.
of the Skull in the Ostrich Tribe.
Phil. Trans, vol. 156, 18G6, p. 113.
78.
Parker,
W. K.
—On
Remnants
the
or Vestiges of
Amphibian and Reptilian Structure found
Proc. Roy. Soc. vol.
in the Skull of Birds, both Carinata; and Ratitse.
79.
Parker,
W.
K.
— On
Zool. Soc. vol.
80.
81.
the Morphology of a Reptilian Bird,
xiii.
xliii.
1888.
Trans.
Opisthocomus cristatus.
1891.
W. N. — Note on the Respiratory Organs oi Rhea. Proc. Zool. Soc. 1883.
Pycraft, W. p. — A Contribution to the Pterylography of the Tinamiformes. Ibis,
Parker,
1895,
p. 1.
82. Pycraft,
W. P.— Contributions
Part
to the Osteology of Birds.
II.
Proc. Zool. Soc. 1898,
pp. 958-989.
83. Pycraft,
W.
P.
Wing.
84.
Rathke, H.
— Some
Facts concerning the so-called " Aquintoeubitalism " of the Bird's
Journ. Linn. Soc, Zool. vol. xsvii. pp. 236-256.
— Anatomisch-philosophische
enbein der Wirbelthiere.
86.
Remouchamps, E.
Remouchamps, E.
87.
RoLLESToN, G.
88.
RoLLESTON, G.
85.
vol.
ii.
— Sur Glande gastrique
— Archives de Biologie,
la
t. i.
—The
90. Sappey, p. C.
Bull. Ac. Belg.
t. 1.
1880,
p.
114.
1880, pp. 583-594.
Collected Scientific Papers and Addresses,
740.
p.
Rothschild,
du Nandou.
Harveian Oration, 1873.
— On
Homologies of certain Muscles connected with Shoulder-Joint.
Linn. Soc. xxvi. (1870),
89.
Untersuch. iiber den Kiemenapparat und das Zung-
Riga, 1832.
W.— The
p.
Genus Aptcryx.
—Recherches sur FAppareil
91. Sclater, p. L.
Trans.
609.
Nov. Zool. vol
vi.
1899, pp. 361-386.
respiratoire de Oiseaux.
1847.
— List of Vertebrated Animals.
92. Stirling, E. C.
and Zeitz, A. H. C.
— Preliminary Notes on Genyornis newtoni, &c.
Tr.
Roy.
Soc. S. Australia, xx. 1896, p. 171.
93. Stirling, E. C. and Zeitz, A. H. C.
newtoni.
94. SusHKix,
Trans. Roy. Soc. S. Australia, xx. 1896,
P. P.
ralistes,
—Description of the Bones of the Leg and Foot of Genyornis
—Zur
Moscou,
95.
Van Bemmelen,
96.
Wedemeyer, G. H.
97.
Wenckebach, V.
J. F.
xvi. liv. 2, 1899, pp.
— Onderzoek van ecu
191.
p.
Nouv. Mem. Soc. Imper. des Natu-
Morphologie des Vogelskelets.
1-163.
jK/jea-embryo.
Tijd.
Ned. Dierk. Ver. 1886-87,
p. ccv.
L. C.
muskulose Beutel.
F.
Leydeu, 1888.
—Der mit der Luftrohre
des
Emu
Casoars in Verbindung stehende
Froriep's Notizen, 1824—25, p. 7.
— De
Outwikkeling en de
Bouw
der
Bursa
Fabricii.
Inaug. Diss.
PHTLOGE^T OF THE PAL^OGNATH.^<: AND NBOGNATH^.
98.
99.
28"
—
Woodward, A. S. Vertebrate Palaeontology, 1898.
Wray, R. S.— On some Points in the Morphology of
the
Wings of
Birds.
Proc. Zool. Soc.
1887, pp. 343-356.
100.
WuNDERLicH,
C.
— Beitrage zur vergleicliendeu
uutereu Kehlkopfes der Vogel.
loi. Zander.
— Uebcr das Gefieder
Kouigsb. xxix. 1889,
Anatomie nnd Eutwickelungsgeschichte
Nov. Act. Ac. Leop.-Carol.
des aiuerikanischen Strausses.
xlviii.
Schrift.
der.
1884.
phys.-okonom. Gesell.
p. 31.
EXPLANATION OF THE PLATES.
PLATE XXII.
Common
Casuarius casunrius.
formerly at Tring,
now
Cassowary
(p.
113).
(Drawn from
life
from a bird
living in Dublin.)
PLATE
Casuarius casuarius beccarii.
a bird formerly at Tring,
now
XXIII.
Beccari's Cassowary (p. 116).
(Drawn fiom
from
life
in the Society's Gardens.)
PLATE XXIV.
Casuariiis casuarius salvadorii.
from a bird formerly
Salvadori's Cassowary (p. 120).
alive at Tring,
now mounted
in the
(Drawn from
Tring Museum.)
life
PLATE XXV.
Casuarius casaarius austraUs.
from a bird that lived ten years
Australian Cassowary
at Tring,
and
is
now
(p. 123).
(Drawn from
lif(>
living in the Society's Gardens.)
PLATE XXVI.
Casuarius casuarius violicoUis.
life
Mauve-nccked Cassowary
(Drawn from
(p. 122).
from the type specimen formerly at Tring, now living in the Society's Gardens.)
PLATE XXVII.
Casuarius casuarius intensus.
Blue-necked Cassowary
(Drawn from
(p. 121).
from the type specimen formerly at Tring, now mounted in the Tring Museum.)
life
PLATE XXVIII.
Casuarius bicarunculafus.
a bird formerly at Tring,
now
Two-wattled Cassowary
(p. 129).
living in the Society's Gardens.)
(Drawn from
life
from
284
.MK. ^V. p.
Pl'CKAFT UN THE iJOEPHOLOGY A>'U
PLATE XXIX.
One-wattled Cassowary
Casiiarms imappendicvlatvs.
(Drawn from
132).
(p.
life
from a bird in the possession of Mr. Blaauvv.)
PLATE XXX.
One- wattled Cassowary
Casuarius unapjyendicuJahif^.
(p.
132).
Whole
(The figure of the adult was made from Mr. Blaauw's bird
and young.
was drawn from
a bird at Tring, the skin of
which
birds, adult
;
preserved in the Tring
is
the young
Museum.)
PLATE XXXI.
Casuarius unappendiculatus occijjHalis.
Jobi
Island
head, neck, and naked parts were drawn from a bird
Cassowary
now
living
(p. 185).
in
the
(The
Society's
Gardens, the casque from the mounted specimen in the Paris Museum.)
PLATE XXXII.
Casuarius unappendiculatus aurantiacus.
Eastern One-wattled Cassowary
from the type now living
in the Berlin Zoological Gardens.)
(Drawn from
life
(p.
136).
PLATE XXXIII.
Casuarius philijn.
now
Sclater's
Cassowary
(Drawn from
(p. 138).
life
from the type
living in the Society's Gardens.)
PLATE XXXIV.
Casuarius
i^o^l^uanus.
Westermann's Cassowary
(p. 139).
(Drawn from
life
from
two specimens formerly at Tring and the Society's Gardens, now preserved in the
Tring Museum.)
PLATE XXXV.
Casuarius 2>(i2niamis edwardsi.
life
from a specimen formerly
(Drawn from
the Tring Museum.)
Milne-Edwards's Cassowary
at Tring,
now mounted
in
(p. 141).
PLATE XXXVI.
Casuarius picticollis.
in Gould's
'
Birds of
New
Painted-necked Cassowary
(p. 143).
(Drawn from the plate
Guinea.')
PLATE XXXVII.
Casuarius ijicticoUis hecM.
Hecks Cassowary
(p.
144).
the type specimen living in the Berlin Zoological Gardens.)
(Drawn from
life
from
—
PHYLOGE.XY OF THE PAL/EOGNATlIJi AND
NEOGNATiLE.
285
PLATE XXXVIII.
Casuariits loricv.
l^oria's
Cassowary
(Drawn from
(p. 142).
a sketch
made from
a fresh-killed bird by Dr. Loriu.)
PLATE XXXIX.
Bennett's Cassowary (p.
Ca.wariiis hennctti.
formerly at Tring,
now
(Drawn from
14-3).
in the Zoological Society's
life
from a bird
Gardens.)
PLATE XL.
Map showing
geographical
the
distribntion of the six
subspecies of Casuariu!<
cuauarius and of Casuarius hicarunculatus.
PLATE
Map
sliowing the geographical
XLI.
distril)ution of the three subspecies of
unapiicndiculatus, two of Casuarius jja^/iiaims, tvvo of Casuurius
Iori(v,
and Casuarius
XLII.
The dorsal aspect of the Skull
1.
The
Casuarius
hennetti.
PLATE
Fig.
jjicticollis,
CatmariuH
(figs.
1-4).
Bhea americana (p. 270), to show the form of the temporal fossa.
fused mesethmoid plate, the form of the lachrymals and nasals,
skull of
the partly
and the broad nasal process of the premaxilla.
Fig. 2.
The
skull of
Lronmus
noMV-hollandiff
(p.
270), showing the rod-like nasal
process of the ])remaxilla, the fused lachrymal, broad interorbital region, and
widely separated temporal
Fig. 3.
fossaj.
Tiie skull of Struthio molyhdophancs
{\i.
269), showing the supra-orbital ossicles
running backwards from the laclirymal to the
process of the premaxilla.
Fig. 4.
The
skull oi
Apteryx
and the absence of
The
niesethnioidal suture
ausfjrilis (p. 269),
its
frontal,
is
showing the small
supra-orbital processes,
and the broad nasal
obliterateil.
size of the
lachrymal
the complete fusion of the
nasal pmcess of the premaxilla, and the rounded form of the frontals, which
do not protect the orbit above.
(figs.
5-8)
showing
the
Ventral aspects
Fig. 5. Tlie
ventral
aspect
of
fig.
J,
processes of the premaxilla and
large
size
of
tlie
palatine
of the maxillo-palatine processes.
The
peculiar relations between the vomer, palatine, and pterygoid (desciibed on
p. 207), and the anterior end of the parasphenoidal rostrum between the
anterior cleft of the vomer.
VOL. XV.
— PART
V.
No, 2o.
Ikccmbcr,
J
900.
2 r
286
MK. W.
Fia;. 6.
Ventral
P.
of
aspect
backwards on
PTCRAFT ON THE MOEPHOLOGT AND
fig.
2,
showing the
to the pterygoid.
The
broad
vomer and
its
continuance
palatines articulate with the external
border of the foot of the vomer and external border of the pterygoid.
Rhea the
between pterygoid and vomer
articulation
is
In
concealed by the
palatine.
Fig. 7. Ventral
aspect of
fig. 3,
showing the
8.
the premaxillse.
The
Ventral aspect of
fig. 4,
which does not now-
Note the absence of the palatine processes of
articulate with the pterygoid.
Fig.
vestigial vomer,
palatine has fused with the external pterygoid border.
showing the complete fusion of the vomer, pterygoid,
and palatine.
Additional
T.etticrs.
= nasal.
= occipital condyle,
/ja. — palatine.
jxi.r. = parasphenoidal rostrum.
jB.jom«. = palatine process of premaxilla.
= paroccipital process.
jo/. = pterygoid.
= supra-orbital bones from lachrymal
= anterior basi-cranial fontanelle.
ch/.A. = antrum.
6.6./>. = basitemporal platform.
b.p. — basipterygoid processes.
Ay ./or. = hypoglossal foramina, 1, 2, 3.
= lachrymal.
= lacln'y mo-nasal pillar.
= lambdoidal ridge.
ff»^6/.
?i.
o.c.
)u./j.
/.
/.;;.
«.
/./•.
= mesethmoid.
mx.p. = maxillo-palatine process.
«./(m>r. = nasal process of premaxilla.
to
frontal.
7H.e.
= vomer.
t7tf^./or. = vagus foramen.
«;.
PLATE
XLIII.
The immature Skull.
Fie
1.
Dorsal aspect of the palatine bones of Aidenjx mantelli
(p.
204), to
show the
sutures, as yet unclosed, between the vomer, pterygoid, and palatine.
Fig. 1 a. Ventral aspect of the same, to
show similar
sutures.
Fig. 2. Sagittal section of skull of Apteryx austraUs mantelli, after Parker, for the
purpose of comparison with iJromu'us.
Note the appearance of the squamosal
on the inside of the skull in Apteryx.
Fig.
3.
Ventral aspect of the skull of
with Bhea.
Note the
Emeus
(p. 270), for
the purpose of comparison
similarity in tlie relations of the pterygoid
Fig. 4. Sagittal section of cranium of
embryo Dromceus novce-hollaudke,
and vomer.
to
show the
separate bones.
Fig. 4
a.
Note how completely the squamosal conceals and
in the nestling Penguin the
the pro- and opisthotic bones.
Outer view of
protects
squamosal
Fig,
0.
is
fig.
4.
deeply emarginate posteriorly and exposes the otic bones.
Ventral aspect of base of skull of lihca americana, to show sutures.
.
PHYLOGENY OF THE PAL^OGNATH.E AND NEOGNATH.E.
Fig. 5 a. Lateral
aspect
vertical
spine
Ekea americana,
of skull of
(lachrymo-nasal
pillar)
from
show
to
the
287
Note the
sutures.
maxillo-palatine process,
which replaces the maxillary process of the nasal the remarkably small
and the outer and inner views of the lower jaw.
of the quadrato-jugal
;
size
;
Fig. bb. Lateral aspect of pterygoid of
Rhea americana
(p. I'Jl), to
show
its
relations
with the vomer.
Fig. 5
Fig. 5 d.
hmer
aspect of mandible
Fig. G. Quadrate of
Bubo
Nat.
head.
Fig. 7.
to
show
sutures.
of tig. 5, to
show
sutures.
Outer aspect of mandible of
c.
fig. 5,
7)iaximus, to
show double, widely separate, squamosal
size.
Quadrate of Dromceus irroratusl, posterior
Nat.
head.
Fig. 8. Quadrate
articular
aspect, to
show
single
articular
size.
of Apteryx oweiii,
posterior
aspect,
to
show double
squamosal
head.
Additional Letters.
= angalare.
=
als.
alisphenoid.
ar. = articulare.
«»/. = antrum of Higlimorc.
b.s. = basisphenoid.
= dentary.
CO/'. = coronoid.
ep.o. = c])iot\c.
e.r. = exoccipital.
a.
(/./<.
= otic
o.r.
=orbital process of quadrate.
head of quadrate,
= parietal.
p.f. = precoudylar fossa.
^/•o. = pro-otic.
n.a. = supra-augular.
A\o = supra-occipital
= spleuial.
/>.
«?.
.
syj.
«(/.=:
squamosal.
= squamosal head of quadrate.
v«(/./. = vagus foramen.
= vomerine process of premaxilla.
= lachrymo-nasal process of premaxilla.
/./. = floccular fossa.
= frontal.
»*. = meatus interuus.
mes. = mesetlimoid.
= opisthotic.
.S(/./«.
//•.
i;o./;/".
/./».
o/j.
PLATE XLIV.
Fig. 1. Dorsal
aspect of the skull of an adult Droma'ns ater (p. 270), to show the
conspicuous, embryonic, cerebral prominence and the fused lachrymals.
Fig. 1 a. Lateral view of the
and temporal
same
skull (fig.
1),
.showing the cerebral
prominence
fossae.
Fig. 2. Dissection of a skull of a
show the form and
young Dromceus novcf-hollandice, lateral view, to
of the mesethmoid for comparison with fig. 3
])ositi(3n
et seq.
Fig. 3
The mesethmoid of
a
young Casuarius unappendiculatus
(p. 199), to
show the
mesethmoid and the large median superposed element forming the centre
the casque of the adult.
of
ME.
288
Fig. 4. Lateral
\V.
p.
PTCEAFT ON THE MOEPHOLOGT AND
aspect of a portion of the skull of Casuarius sclateri, showing the
frontal, nasal,
and median elements of the casque.
Fig. 4
f/.
Dorsal aspect of
Fig. 4
h.
Dissection of a skull of Casuarius sclateri^=fi.g. 3
fig.
3 a.
a,
showing the mesethmoid
and a portion of the ventral border of the median ossification of the
casque interpolated between the mesial border of the anterior ends of the
frontal.
young Casuarius unapjyendiculatus after
removal of the right innominate, to show the great height of the neural
Fig. 5. Lateral view of the synsacrum
of a
spines of the verlcbrfe.
Fig.
Lateral view of the synsacrura of an
6.
prepared for comparison with
embryo Drommis
fig.
Note that
5.
novce-hollandia; (p. 212),
this stage the neural
at
spines have not yet risen above the centra of the vertebrae, but form only a
series of lateral wings.
Fig. 6 a. Dorsal aspect of a portion of the synsacrum
that
the
condition
Fig. 6
neural
do not
above the
rise
G), to
show more
centra
adult
the
in
as
clearly
(fig. 5).
Lateral view of
b.
spines
(fig.
the same pelvis
(fig.
6),
to
show the separate elements
of the innominate.
Additional Letters.
= anti-trochanter.
/.c. = troutal moiety of casque.
o.yy./r.
a/ii.
process of frontal.
= pectineal process.
= pubo-iscbiadic fissure.
s.c. = sacral vertebrae 1-2.
]
= ilium.
il.
= orbital
/>.=pubis.
/).;3.
= ilio-iscliiaclic foramen.
= ischium.
l-6 = himbar vertebne 1-G.
lb. l-2 = lumbo-sacral vertebrse 1-2.
»ne.c. = median element of casque.
?(.c. = nasal element of casque.
o6. = obturator fissure.
jmbo-isch.f.
?7.w./.
js.
s.cd.
II).
= sacro-caudal
vertebrae.
= sacro-thoracic
=
vestigial thoracic
v.
«.»ie. = ventral border of
,
^
'
,
vertebrae.
.
s.M.
}
,,
rib.
mesethmoid.
PLATE XLV
Fig. 1.
The buccal
cavity
of Eltea aiiiericana (p. 248),
to
show the large choan^
characteristic of the Falceofjnatlim.
Fig. 2.
A
neossoptile of Calodromas elcgans (p. 162), after Pycraft, to
aftershaft, equalling the
shaft,
main
shaft in size, the
and their production forwards into horny filaments resembling the
grooved ribbon-shaped bands of the young Ostrich
Fig. 2 «.
A
show the large
complex rami of the main
radius from the distal end of the
main
shaft.
(fig. 7).
—
PITYLOOEW OF THE PAL.EOnXATILE
Pig.
A
3.
A\|) XEOOXA't'llE.
neossoptile of lilim amrrirnnn (p. TG4): neither
have a distinct
main nor
tlic
289
Llie aflersliaft
The main shaft is represeuted by three stiffened
Tinamnns and Ostrich, the aftersluift by
rliachis.
rami, prnduced into filaments as in
nnmerous slender rami.
A
Fig. 3 a.
A
Fig. 4.
portion of a radius from
neossoptile of Dronnt'titi iiora'-JinUaiidiii- (p.
distinguishable
A
Fig. 4 a.
of a radius
jKirtinn
suppression of the
A
Fig. G.
poitidu of a
A
radius of
neossoptile of
tStnitlilo
rami described on
Fig. la.
Note
G.
fig.
p.
irnneJiis,
A
fig.
portiou of a radius of
Fig. 8. Dissection
There
that
is
further
4 a
aftershaft.
numerous and
are
fila
figs.
no
and
These arc flattened
7.
of the air-sacs of the
left
large,
5.
as in (^'nji>tnri (fig. 2).
of a (kisiiarhis intajipeniJiciilatiis
side
showing the three divisions of the anterior intermediate sacs and the
Note the position of the anterior end of
positions of the ostia throughout.
the posteri(;r abdominal air-sac
mediate.
more or
The
less
intermediate.
9.
still
" eye-spots."
Note the
Fig.
3-7.
104, and representing the main shaft, and the numorons
the aftershaft.
a
barely
is
showing the three peculiarly modified
downy rami representing
(p. 252),
figs.
showing the
casiiarins,
('r/siKin'n.s
indicating a less degenerate condition than
Fig. 7.
aftershaft
fila.
neossoptile oi yljjfef//x nioufclli (p. 103).
A
Fig. G a.
of
'i'lie
compare with
:
fila.
Fila are yet represented but arc small.
4.
fig.
aliscnce of
tlic
l^\'l).
the main shaft well developed
;
portion of a radius of
A
Fig. 5.
Note
3 (p. 104).
fig.
posterior abdominal, furthermore,
seen to take the form of
of the kidney.
It encloses a part
lung of
the costo-]iulmonary
muscle,
of the
ramifications.
is
sausage-shaped sac adherent to the inner wall of the posterior
left
Dissection
— forcing down the roof of the posterior inter-
252), to
show
aud euto-bronchia and
their
C<tsif(fr/us uiiajjjicrid/atlafiis
meso-,
ecto-,
Note that the ento-bronchia
1
and
3
(p.
between them supply
the tubulae of the ventral surface of the lung described on p.
systems fed by ento-br(jnchium
ento-bronchiiim 2
lie
1
252.
all
The
have been exposed by dissection, those fed by
caudad of these and are concealed by the aponeurosis.
Fig. 10. Ventral aspect of a dissection of the
wing of an adult
Cus/iariiis vasuariKS,
show the hypertrophied calami described on p. 1C5.
Dorsal aspect of a preparation of the wing of an
to
Fig. 11.
Ostrich.
After
Wraij.
Fig. 12. Dorsal aspect of a portion of the
wing of
Jllica
americauu
(p. 150), to
show
the remarkable distribution of the coverts in transversely oblique bundles.
Fig. 13. Ventral aspect of the
wing of Aptenjx mantclli
(p.
of remigesand their remarkable resemblance to the
on
VOL. XV.
IGl), to
show the number
young Cassowary described
p. 105.
— PART
V.
No. 24.
Ikcemher, 1900.
2
s
OX THE MORPHOLOGY
290
A
Fig. 14.
ETC.
remex of a nestling
OF THE PAL.^OGNATH^ AND NEOGNATH^.
Casiiarius aiisiralis, dorsal aspect, before the loss
of
the rliachis described on p. 165.
show the peculiar rhachis.
the rhachis of figs. 14-14 a (p. 165).
Fig. 14
ff.
Ventral aspect of
Fig. 14
h.
Section of
Fig. 15.
A
fig.
14, to
reme.\ of Apteryx mantelli (p. 166), for comparison with that of the
Cassowary described
p.
young
165.
Additional Letters.
= aftershaft.
f/.i?2/.l-3 = anterior intermediate
ff.s.
=ala
air-sacs 1-3.
spuria.
para.b.--= parabi'oncliia.
«y>.= aponeurosis.
c.
<".c.
= calamus.
= carpal covert.
p.r.
rh.
= costo-pulmonary muscle.
= carpal remex.
= cubital remiges.
* = hypothetical carpal covert.
r.m.
c.p.m.
c.?'.
„
„
= ecto-brouchia.
ent.
=ento-bronchia.
= eustachian
/. = filum.
A. = lieart.
c?<,
remex.
,,
aperture.
,,
= passage
= kidney.
?«i.
= traverses
=meso-bronchium.
= metacarpo-digital remiges.
0. = ostium posterius.
o6/..se/?. = oblique septa.
0. ent. 3 = oi't'\a of cnto-bronchium 3.
= ostia of outer and inferior
o.mb.
l-.'2
= passes
7>.6.
„
1
to
„
= pre-bronchial
„
1
to
ento-
vestibule.
lateral
pre-bronchial sac.
whole
bronchium
sub-bronchial sac.
air-sac.
length
1 to vestibule.
from ento-bronchium
= traverses
[
= ost\& leading into
entire
bronchium
branches (1-2) of meso-bronchium.
o.p.b.s.=
from ento-bronchium
vestibule.
j)».rf.r.
o.syb.
rami.
t.
= eye-spots.
e<?/.
= ramus.
= ribbon-shaped prolongations of
r.g. = rima glottidis.
= tougue.
t-f. = terminal filaments of the rami.
t.mj. = tectrices majores.
t.md. =
medise.
„
i.m. =
minores.
t.ms. =
marginales.
r.r.
€.?-.
A.
vein.
= rhachis.
i-h." = rhachis of aftershaft.
= choaute.
c/. = claw.
e.
= pulmonary
rd..=:radins.
cA.
**=
= posterior intermediate air-sac.
p.ahd. = posterior abdominal air-sac.
p.a. = pulmonary artery.
pant.
fl.
i
~
I
length
ento-
4.
traverse lateral meso-bronchia into
atiteri(U'
and
posterior
mediate air-sacs.
inter-
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