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Some seven years later Owen (1866) elaborated on his description after
receiving additional skull fragments from Darling Downs which were sent
to him by Edward Hill of Sydney. When pieced back together, these
fragments made a much more complete specimen than the skull from lake Colongulac.
The alveoli of the maxillary's pre-carnassial teeth were all present and
the canine itself was still in position. Owen then determined that
Thylacoleo
must have had an enormous tusk-like I1/ and two smaller
incisors on the premaxilla while the maxilla must have carried the small,
blunt canine followed by one or two small premolars positioned medially
to the anterior root of the carnassial. He also made mention of the
double wear facet on the medially-directed occlusal surface of P3/
which
shows a slicing action was had occurred against both P/3
and M/1. Owen also pointed out the fact that
the glenoid cavity was not semicylindrical in design as in typical placental
carnivores. Though it is transversely elongate and bears processes
limiting lateral excursion of the condyle, its articulating surface is
slightly convex ventrally and extends forward for about 2 cm from the postglenoid
process.
Information on the mandible
was provided through a photo and a drawing of Wellington Caves material
sent by Mr. Gerard Krefft, Curator at the Australian Museum, Sydney.
From these, Owen demonstrated that there was only one large tooth at the
anterior perimeter of the dentary, though its morphology could not be determined
as the crown was no longer present. Only two small alveoli lay between
this tooth and the mandibular carnassial. Based upon this pictorial
information, Owen presumed that the thylacoleonid taxonomic relationship
lay with the diprotodontid marsupials as opposed to the carnivorous dasyurids.
Owen gave Thylacoleo's dental formula as I 3-3 / ?, C 1 / ?, P 2-2
/ 2-2, M 1-1 / 2-2 = 24 denoting his incertitude as to whether a second
incisor or a canine lay behind I/1 and considering
that the small anterior premolar was double rooted. Owen restated
his belief in the carnivorous nature of Thylacoleo, and recorded
that it "exemplifies the simplest and most effective dental machinery for
predatory life and carnivorous diet known in the mammalian class, It is
the extreme modification, to this end, of the Diprotodont type of Marsupialia"
(Owen 1866: 81).
More fossil material
from Darling Downs had reached Owen by 1871 and from this he described
I1/, C1/, the two small premolars
anterior to the maxillary carnassial and a mandibular incisor in which
the base of the crown was still present. However, the premaxillary-maxillary
suture was not well defined and appeared to run through the alveolus of
I3/ which Owen had therefore labeled as C1/,
while the actual canine became P1/. The photos
provided by Krefft had served as a basis for illustrations of isolated
teeth in this paper and, in the next year, Krefft (1872b) cited that his
images of I2/, I3/, and C1/
had been labeled P1/, C1/, and
P3/ respectively by Owen. This mistake was soon
made clear. In 1868, Flower had published a drawing of the
skull in which the canine was correctly labeled and in 1876 McCoy provided
a comprehensive description of the path of the premaxillary-maxillary suture
in new fossil specimens from Lake Colongulac. The suture was demonstrated
to approach the alveolus of I3/ but then, at the periphery
of the jaw, it was redirected back to run through the alveolus of the canine.
The serrated enamel
on the lateral margin of each I/1 was noted by both
Owen (1871) and Krefft (1872b). Krefft also pointed out that the
enamel was restricted to the tips of the crowns on the interdental surfaces
of these particular teeth. He also noted wear facets in this region
which indicate that there was movement between the mandibular incisors
and that, thus, the mandibular symphysis could not have been fused.
Each of the large maxillary first incisors were shown to bear a wear facet
on its lingual surface illustrating that the mandibular incisors passed
behind the maxillary ones upon jaw closure. The crowns of I2/
and I3/ also exhibited evidence of abrasion at their
tips as a result of contact with I/1.
The next notable breakthrough
in the study of Thylacoleo carnifex came about in 1883 when Owen
(1883a) ardently acknowledged receipt of a maxilla and a mandible from
the Wellington Valley, both of which still retained all of their teeth.
The mandible possessed an intact, transversely-elongated condyle and a
deeply inflected mandibular angle but the coronoid process, unfortunately,
was broken. The broad, anteriorly-directed profile of the latter
was finally established in 1887 from a cast of a complete dentary forwarded
to Owen to G. P. Ramsay, successor to Krefft. In this, Owen's final
compendiary publication regarding
Thylacoleo, he reprised his conjecture
that the animal was a diprotodontid which had become adapted for a life
of carnivory.
| The
orbit (eye socket) of Thylacoleo's skull. The arrow points to the
postorbital bar. Among marsupials, this feature known to exist only
in Thylacoleo and Thylacosmilus. In this particular
specimen, the centre of the postorbital bar is missing. In an intact
skull, the orbit would be completely surrounded by a ring of bone. |
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Of the
skull anatomy discussed thus far, no mention has yet been made of two nearly
unique features of
Thylacoleo. The orbit of this animal is
delineated posteriorly by a slender bar that is missing from most fossil
specimens. This postorbital section of bone is also found in the
extinct Thylacosmilus of South America, but is not present in other
marsupials. It was illustrated without comment in a reconstruction
of the skull in Lydekker's catalouge (1887, fig. 28) but notice was drawn
to this peculiar characteristic by Anderson (1929) and by Gill (1954).
The second feature of note, one which is totally unique to Thylacoleo,
was also remarked upon by Anderson. Thylacoleo's sphenopalatine
foramen, through which the orbit and nasal cavity are in contact, is significantly
larger, relatively, than that seen in any other land mammal. No explanation
for why this should be has been put forward. In 1956, Woods produced
an in depth description of the skull of Thylacoleo based upon the
Darling Downs specimens held at the Queensland Museum. From one skull
he was able to create an endocranial cast which he compared with the one
described by Gervais as far back as 1869. |
Among the marsupials,
the brain of Thylacoleo would appear to most closely resemble that
of the phascolomids. The ear region (including the stapes) as well
as the temporomandibular joint, however, show affinities to those of the
phascolarctids. Woods also pointed out the fact that no deciduous
teeth had been found in any specimen.
Presently, skull and
dental material of Thylacoleo carnifex have been found in every
state in Australia. The Western Australian and Tasmanian specimens
are few in number as compared to that discovered in the other states, but
it would seem that they belonged to animals that were of smaller build
than those hailing from the eastern states. This discrepancy in size
has been examined by Finch and Freedman (1982).
POSTCRANIAL MATERIAL:
A metacarpal was described
by Owen (1859), a radius, ulna and terminal phalanx (1883a) and a pelvic
girdle (1883b) while De Vis made a description of a femur (1887) and a
radius (1900) and Longman (1925) classified a calacaneum to
Thylacoleo.
Ungual phalanges were discovered by Krefft, one of which he described (1870a)
as Mylodon australis although Owen (1871, 1877) was inclined
to consider them as being Thylacoleonine. Although many of these
specimens had been recovered from the same localities as Thylacoleo
skulls, however, there was no solid evidence that they were genuinely parts
of a Thylacoleo.
The sole published record
of postcranial material which was undoubtedly referable to Thylacoleo
was that of Wells and Nichol (1977). They described the Thylacoleo
manus in both articulated and disassembled manner and were of the belief
that the forelimb stance was digitigrade. According to these authors
considerable mobility would have been exhibited by the pollex and appeared
to be pseudo-opposable to the pisiform. The pes was not complete
but its structure inferred a plantigrade posture and a hallux that was
divergent from the other digits.
THE MOREE THYLACOLEO:
| A nearly
complete skeleton of Thylacoleo was
discovered in a loam pit in 1966 near Moree in northern New South Wales.
The right scapula and right ilium were still in association with the vertebrae
and the vertebral groups articulated together to form a complete column
apart from the terminal caudal units. The limb bones were broken
but could be pieced back together to make a complete forelimb and hindlimb
which were missing only the pes. The adult specimen was in association
with a juvenile, represented only by a large skull fragment and a section
of the lower jaw. Found with these was the crushed skull of a Thylacoleo
pouch-young. This material has been used in examinations of mastication,
feeding habits and locomotion (Finch and Freedman 1982, Finch 1981). |
| The
Moree Thylacoleo, one of the most complete skeletons ever found
of T. carnifex. Unearthed at a loam pit in New South Wales
in 1966, it offered the first solid glimpse of Thylacoleo's postcranial
anatomy. |
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