| THYLACOLEONID
MATERIAL CLASSIFIED TO DIFFERENT GENERA AND SPECIES:
Following is an account
of genera and species which have been described at various times since
1859 and studies by Archer and Dawson (1982) have defined the position
of some of these taxa.
Plectodon
Krefft, 1870: A brief description was published by Krefft of material
which included some badly damaged mandibular incisors said to come from
an animal "allied to Thylacoleo" which he named Plectodon.
No type specimen was named and no details regarding its locality of discovery
were mentioned. A specimen was sent to Owen but no species name was
ever recorded, despite the fact that Krefft made further references to
Plectodon
in later publications (1870b, 1882) and provided an illustration of the
incisor (1882). Plectodon is considered by Archer and Dawson
(1982) as a junior synonym of Thylacoleo Gervais.
Mylodon? australis
Krefft, 1870: Krefft found a large, hooded ungual phalanx amongst
some of the material collected from the Wellington Caves. It rather
resembled that of a South American ground sloth, Mylodon.
He assigned the specimen to the genus Mylodon? and made it the type
of Mylodon? australis. A short description was published (Krefft
1870a) however, it was Owen (1871 1877) who first illustrated the specimen
from a series of photographs of two variously shaped types of phalanges
which had been sent to him by Krefft. Owen produced figures of both
types of phalanges and, by excluding all other marsupials known to have
existed in the discovery locality at that time as being of the improper
size, he expressed that Thylacoleo must have been the possessor
of the curved ungual phalanges. He rejected the possibility that
Mylodon
had ever existed in Australia. Etheridge (1918) examined the confusing
situation which had developed out of Owen's referral of two distinct types
of phalanges to Thylacoleo and the listing by Lydekker (1887) of
a cast of a third form of phalanx as also being ascribed as that of a thylacoleonid.
Krefft's hooded, ungual bone was likely referable to Thylacoleo.
Of the two unhooded phalanges, the highly arched and compressed type was
morphologically most similar to that of the great flying phalanger while
the more elongate, less compressed claws were more like those of the koala.
In a study (1982) Archer and Dawson demonstrated that Thylacoleo
carnifex is a senior synonym of Mylodon australis.
Thylacoleo oweni
McCoy, 1876: The innate difficulties in having to base the taxonomy
of fossil species upon a few incomplete specimens are highlighted by the
studies of McCoy (1876). He had been given a fragmentary skull and
mandible in 1875 collected by Adeney from the same locality at Lake Colongulac
as the specimens that had been sent to Owen thirty years earlier.
The skull appeared to be lacking part of Owen's specimen and bore all precarnassial
teeth in situ. McCoy made thorough comparison between the
the Victorian material (including the newly obtained mandible) and that
from darling Downs and Wellington Caves (Owen 1859, 1866, 1871) which led
him to differentiate the New South Wales Thylacoleo as a distinct
species, T. oweni.
The basis provided by
McCoy for the new species was as follows:
| a) |
The longer convex margin
of I3/ was posteriorly positioned in
T. carnifex
but oriented anteriorly in T. oweni. Owen had based his description
of T. carnifex, however, upon isolated teeth and had obviously reversed
I3/. |
| b) |
The premaxillary-maxillary
suture traversed the alveolus for the canine tooth in T. carnifex
but was said to meet the alveolus anterior to that of the canine (i.e.
the tooth initially classified as I3/) in Owen's 1871
account. |
| c) |
The dorsoventral depth
of the mandibular ramus was measured anteriorly to the carnassial and again
posteriorly of M/2. The Victorian material showed
the anterior distance to be greater while the T. oweni mandibles
were deeper at the posterior end. |
| d) |
The T. carnifex
palate was narrower across the premaxillary area and 6 mm greater in length
anteroposteriorly that that seen in T. oweni. |
| e) |
I2/,
I3/, and C1/ were larger in the
Victorian than in the New South Wales specimens. Among these alleged
differences, the first two (a & b) have been demonstrated to be invalid
by Owen (1877) whist points c, d, & e would appear to be within the
acceptable variability range shown by T. carnifex. Thylacoleo
oweni is regarded by Archer and Dawson (1982) as a junior synonym of
Thylacoleo
carnifex. |
Thylacoleo robustus
Krefft, 1872: The name of Thylacoleo robustus was given by
Krefft (1872a) to a right maxillary canine found at the Wellington Caves.
His only reason for assigning it to a distinct species was because the
homologous tooth of T. carnifex was said to be reasonably smaller
that that of the new species. T. robustus is now considered
a junior synonym of T. carnifex (Archer and Dawson 1982).
Thylacopardus australis
Owen 1888: An array of fossil specimens from a newly found cave in
Wellington valley, New South Wales, gave Owen a skull lacking a mandible,
and that was smaller than his previous specimens of Thylacoleo.
He called it Thylacopardus australis, the name first being published
in Nature (1888). Owen presented his complete description of the
specimen to the Royal Society of London in 1888, however, the paper itself
was only published by title (1889 or 1888). Two characteristics were
used to make a distinction between this new genus and Thylacoleo,
its small size and its possession of a small molar or 'molariform prominence"
on the posterior margin of the palate behind M1/.
Owen wrote in his yet
to be published manuscript (quoted by C. Anderson in 1929) that "the dentinal
structure" of the molariform prominence has been shown, thus exhibiting
the existence of a second maxillary molar. The manuscript and type
specimen (which was then in the collections of the Geological and Mining
Museum, Sydney) was examined by Anderson and he came to the determination
(1929) that the "M2/" was merely the broken edge of
the post palatine bar and showed no evidence of dentine material.
He held that the animal was in fact Thylacoleo carnifex but could
only have been a young individual on account of its small size and the
fact that it had unworn teeth. Anderson pointed out that even though
Owen's measurements of the skull bones of Thylacopardus and Thylacoleo
displayed a reasonable degree of difference in size, the lengths of the
maxillary carnassials were virtually the same. Therefore, it is unlikely
that Thylacopardus represents a valid genus and species. It
is considered as a junior synonym of T. carnifex by Archer and Dawson
(1982).
.
This map shows the various
localities around Australia where thylacoleonid fossils have been found.
(After
Archer, Clayton and Hand 1984; Vertebrate Zoogeography and Evolution
in Australasia,
Hesperian
Press, Carlisle, Aust.)
Thylacoleo
crassidentatus Bartholomai, 1962: Included in the material
from the Darling Downs area kept at the Queensland Museum is a collection
which comes from Chinchilla, northwestern Darling Downs. These specimens
were thought by Woods (1956) to be significantly different to warrant continuing
study and proposed that they were of Pliocene age. All known Thylacoleo
carnifex material is considered to be of Pleistocene age. Acquiring
a partial right mandibular ramus from Chinchilla gave Bartholomai (1962)
the necessary material for the classification of a new species, T. crassidentatus.
Bartholomai lists the basic distinguishing characteristics of this species
as compared to T. carnifex as:
| a) |
P/3
is broader posteriorly and more convex buccally than is that of T. carnifex. |
| b) |
M/1
is wider over the anterior and posterior roots than as seen in T. carnifex.
The posterior section of the crown possesses a wide median ridged area
and a shallow posterolabial fossette. M2 is
also more robust in form and one specimen gives an indication of an alveolus
for M3. |
| c) |
T. crassidentatus'
mandibular ramus is broader and not as concave medially. |
| d) |
I/1
is more recumbent, with the angle between the tooth and the base of the
mandible being smaller. The maxillary material was fragmented, and
did not display and notable distinctions from T. carnifex although
one specimen may have had an M2. |
Further material which can be referred
to T. crassidentatus has been described from the Allingham Formation
(Lower Pliocene) of Northern Queensland (Archer and Wade 1976) and at Bow
in New South Wales (Archer and Dawson 1982).
Balladonia specimens:
Glauert (1912) and Merrilees (1968) both refer to Thylacoleo from
caves near Balladonia, Western Australia. Merrilees makes note that
Glauert failed to differentiate between the various caves around Balladonia
but Merillees himself mentions Wonberna Cave as being the source of Thylacoleo
"probably not carnifex". This animal was significantly smaller
than T. carnifex and may have been a distinct species (Finch and
Freedman 1982). The age of the deposit from which the material came
is not known.
Thylacoleo
hilli Pledge, 1977: An isolated left maxillary carnassial
of Thylacoleo found in 1956 in a cave at Curramulka, South Australia
has been described by Pledge (1977). The fossil is of unknown age
but is thought to be Late Miocene or Early Pliocene. It is quite
peculiar due to its small size (Finch and Freedman (1982), being roughly
half the length of the P3/ of T. carnifex.
Because of this, it has been designated a new species, Thylacoleo hilli.
A discovery of new material referable to this species at Bow, New South
Wales has been reported by Archer and Dawson (1982).
Wakaleo:
From the Miocene deposits of South Australia and the Northern Territory
have come thylacoleonids which are considered by Clemens and Plane (1974)
to be a genus which bears notable dissimilarities from Thylacoleo.
These animals had a different dentition, as they had retained three mandibular
molars, had a comparatively shorter carnassial tooth and bore only one
precarnassial premolar. The latter is a characteristic not evident
in Thylacoleo species. Such dissimilarities, they gathered,
was a sufficient basis for constituting a new genus, Wakaleo.
The species name, W.
oldfieldi, was given to the South Australian specimen from lake
Ngapakaldi specimen which consists of a left dentary which includes the
incisor, P/3 and M/1. Also
present are alveoli for P/1 (or P/2)
M/2 and M/3. Because of
wear facets formed on an isolated M/2, it was presumed
that there were two maxillary molars which were positioned posterior to,
and not lingual to, the carnassial.
The fossil from Bullock
Creek, Northern Territory was a fragment of mandible bearing a single molar,
M/1, alveoli for M/2 and M/3,
and a partial alveolus for I/1. The dentary
was somewhat larger than that of W. oldfieldi, and when one takes
into consideration the spatial separation as well as a possible time difference
between the two specimens, Clemens and Plane (1974) felt well-founded in
designating a second species, W.
vanderleuri.
A third species of this
genus, W.
alcootaensis, from the Miocene Alcoota local fauna of the Northern
Territory (Archer and Rich 1982), would appear to be the largest member
of the genus yet found. This specimen, a fragment of a skull, retained
tribosphenic molars, a characteristic which implies secondary modification
of the ancestral quadritubercular molars for a life of carnivory.
Mention is made by Clemens
and Plane (1974) of an edentulous (missing its teeth) skull fragment from
the mid-Miocene Ngapakaldi Fauna. Possibly, it was a thylacoleonid,
as it displays alveoli for an enlarged P3/ and four
maxillary molars. |