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Page 6 ACKMA Journal No. 97 December 2014
CATCHING the MARSUPIAL ‘LION’ by the TAIL:
THYLACOLEO CARNIFEX and the NARACOORTE CAVES
Michael Curry, Liz Reed1,2 and Steve Bourne3
1School of Physical Sciences, The University of Adelaide, Adelaide, SA, Australia; 2School of Physical Sciences, Flinders
University, Bedford Park, SA, Australia; 3Naracoorte Lucindale Council, Naracoorte, SA, Australia.
Introduction
Of all the extinct Australian Pleistocene megafauna
species, Thylacoleo carnifex (the marsupial ‘lion’) has
captured the imagination and interest of people more
than any other. Perhaps it is the allure of its predatory
habits, (Australia’s Pleistocene answer to T. rex); or the
intriguing notion that it used caves as dens (Lundelius,
1966 ). It is certainly an enigma and, as Owen (1866)
suggested, an extreme and meat-eating version of the
otherwise herbivorous diprotodont marsupials.
Spectacular fossil finds over the past few decades have
put to rest much of the speculation regarding its habits
and morphology. Fossil remains found in caves at
Naracoorte in South Australia have played a central role
in solving the puzzle of Thylacoleo. Likewise, this iconic
marsupial has been integral to the history, science,
tourism and interpretation of the Naracoorte Caves. The
aim of this paper is to explore the discovery of
Thylacoleo carnifex at Naracoorte Caves and how this
has influenced the scientific and social history of the
park.
Discovery
The first recorded fossil material of Thylacoleo carnifex
was recovered by Major Thomas Mitchell at Wellington
Caves in New South Wales in the 1830s (Figure 1). It
consisted of only a few isolated teeth and was not
immediately identified (Gill, 1954). Lake Colongulac,
near Camperdown, in Western Victoria, some 150km
east of Naracoorte, bore the first remains to be formally
described by science. The eminent palaeontologist, Sir
Richard Owen, received the fossils in 1855 and
described the bones (parts of the cranium), in 1859. He
interpreted the remains as those of a great marsupial
carnivore and was sufficiently impressed by its
carnassial-like premolars to assert that “…it was one of
the fellest and most destructive of predatory
beasts.” (Owen 1859). Almost at once, controversy
ensued.
William Sharp Macleay, an entomologist with a keen
interest in Australian Natural History, lashed out in the
Sydney Morning Herald convinced of Owen’s folly,
defending Thylacoleo as “A very gentle beast, and of
good conscience” (Macleay 1859). Macleay based his
argument on Thylacoleo’s relationship with other
Diprotodont marsupials, most of which are herbivores.
Gerard Krefft, Curator of the Australian Museum, was
almost equally as unimpressed with Thylacoleo’s
carnivory, opining that it “…was not much more
carnivorous than the Phalangers (possums) of present
time.” (Krefft, 1866). Owen, meanwhile, had received an
almost complete skull from the Darling Downs, in
Queensland and published a more detailed paper,
further describing the skull and teeth of Thylacoleo,
acknowledging its diprotodont affiliation but more
convinced than ever of its carnivorous habits (Owen,
1866). The debate raged back and forth in both the
popular media and scientific literature for some years
(for example see Broom 1898). While there was not
enough evidence to settle the debate, Thylacoleo had
captured the imagination of the public and scientific
community.
The controversy did not end with Thylacoleo’s diet.
Krefft further proposed that an unusually large, clawed
distal phalanx found at Wellington Caves belonged to an
Australian Megatherium (giant sloth), and proposed the
name Mylodon australis (Krefft 1870 - republished in
Mahoney and Ride, 1975 page 198). Owen, who received
photographs of the ungual from Krefft, referred the claw
to Thylacoleo, almost purely by the deduction that it
belonged to a carnivore, stating there was “No evidence
of a Megatheroid…in Australia” and that “there is no
other associated Carnivore corresponding in size…save
the Thylacoleo” (Owen 1871; Figure 2)). Owen later
received and described (again from Wellington Caves), a
mandible, radius, ulna and a similar distal phalanx
(Owen, 1883a) as well as a partial pelvis (Owen, 1883b),
which he attributed to Thylacoleo. However, as an
articulated skeleton was yet to be found, there was still
a question whether these elements were from Thylacoleo
at all. In 1888, Owen presented to the Royal Society a
new Genus in the Thylacoleonidae, although the paper
was never published (Anderson, 1929). He had received
a skull (much smaller than those recovered earlier),
from a newly found cave near Wellington. He named the
species Thylacopardus australis, an allusion to the
animal’s size being more leopard-like, than lion-like.
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“Thylacoleo 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)
Page 7 ACKMA Journal No. 97 December 2014
Early Discoveries at Naracoorte
Father Julian Tenison-Woods was the first person to
describe any fossil material from Naracoorte Caves;
however, he did not report any material attributable to
megafauna species (Woods, 1858; Reed, 2012; Reed &
Bourne, 2013). There can be little doubt he yearned to
find megafauna animals, as he devoted an appendix in
his 1862 book to the fossil discoveries of Wellington
Caves (Woods, 1862). While, he had some success in
finding isolated specimens in other sites around the
southeast of South Australia, he never found them at
Naracoorte. What Woods couldn’t know was that the
remains he sought lay as little as one metre below his
feet as he explored Blanche Cave (Reed and Bourne,
2013).
William Reddan (caretaker of Naracoorte Caves from
1886 to 1919) was the first to discover Thylacoleo fossil
remains at the caves. In Specimen Cave, he found skull
and jaw bones of an unusual animal with “large tusks
protruding” (Anonymous 1908a,b). He forwarded the
bones to his department and they were subsequently
presented to Dr Edward Stirling, Director of the South
Australian Museum (SAM). Stirling identified the
remains as belonging to Thylacoleo and was keen to visit
the site to determine if bones of the post-cranial skeleton
were present in the cave. Fifty years after Owen’s papers,
very few bones of the skeleton were yet known to science,
so the opportunity to expand knowledge of the species
was too good for Stirling to pass up. He decided to do a
systematic search for fossils and arrived at the caves on
Monday 14th
of December 1908, accompanied by his
assistant, Fritz Zietz. They began their search the next
day and found numerous bones, which were taken back
to the museum for further study (Figures 3 and 4).
Stirling later reported on the finds in reports to the
museum board (Stirling, 1908, 1912). Reddan made
some further finds in the newly discovered Alexandra
Cave and these bones were also forwarded to the
museum. Unfortunately Stirling never published the
material.
Renewed interest in the science of Naracoorte Caves
came in the 1950s, with the formation of the Cave
Exploration Group of South Australia (CEGSA). Members
of the group continued systematically exploring, mapping
and surveying the caves. During these surveys, fossil
material (including Thylacoleo) was discovered scattered
throughout the system, adding numerous specimens to
Naracoorte’s collection; but in 1956, an amazing cave
deposit was about to come to light in a somewhat
unconventional way.
The James’ Quarry Thylacoleo
During quarrying operations, Mr Amos James, the
proprietor of Naracoorte Quarry, opened up a previously
unknown cave (5U29) during blasting operations. Within
a small cave he discovered an unusual skeleton and
reported the find to staff at the SAM, who dispatched
Norman Tindale (Curator of Anthropology) and fossil
preparator, Paul Lawson, to investigate. Tindale, an
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Figure 1 - Thylacoleo incisors collected by Mitchell and
sent to Owen at the Natural History Museum, London
Photo: Steve Bourne
Figure 2 - The type of “Thylacoleo oweni”
in the Natural History Museum, London.
Photo: Steve Bourne
Page 8 ACKMA Journal No. 97 December 2014
accomplished anthropologist and entomologist, had
previously published papers on the nearby Tantanoola
Caves (Tindale, 1933) and excavated megafauna fossils
at Rocky River on Kangaroo Island (Hall et al, 1935); a
site that was later to bear Thylacoleo fossils of its own.
Law s o n w as als o e x p erien c e d w ith Ple i s t ocen e
megafauna, having recently accompanied the American
palaeontologist Ruben Stirton to Lake Callabonna in the
state’s northeast, where they famously excavated large
numbers of Diprotodon fossils embedded on the lake’s
shore (Anonymous, 1953).
Unfortunately, Tindale and Lawson arrived a little too
late. With the best intentions, Mr James had covered the
well preserved skull of the animal with a hessian bag to
protect it from the weather. They arrived to find the skull
in pieces, crushed under the weight of the rain soaked
bag, scattered amongst the talus at the base of the
quarry face (Daily, 1960). What remained in the cave was
the well-preserved post-cranial skeleton of Thylacoleo
carnifex, which Lawson painstakingly retrieved from the
sandy sediment of the alcove while balanced on a
wooden ladder some 4 metres above the quarry floor.
The post-cranial skeleton of Thylacoleo, which had
remained elusive since description of the skull by Owen
almost a century before, had finally been discovered.
Previous attribution of post-cranial material to
Thylacoleo by Owen was now confirmed. It was the start
of a rewarding few years for palaeontologists. CEGSA
members found the skull of a juvenile Thylacoleo, along
with some skeletal elements (SAM P12902, P12911), in
Cathedral Cave (5U12), which is now part of the
Naracoorte World Heritage area. Ruben Stirton and
Richard Tedford found another partial skeleton near
Lake Menindee in NSW (Daily, 1960). In 1959, blasting
at James’ Quarry opened a second cave, in close
proximity to 5U29. Another partial skeleton, comprising
most of the front half of the animal and including a
complete – albeit disarticulated - left manus, and
complete forelimb, was added to the museum’s collection
(SAM P2910, Figure 5). Stirton and Tedford took the
material back to UC Berkeley with the intention of
describing the post-cranial skeleton. Unfortunately this
did not eventuate and Gill (1973) suggests that the task
was passed on.
Curiously, despite the excitement of the long awaited
finds, research on the post-cranial skeleton was not to
be published for almost two decades. The material and
inspiration for this research was to come not from the
James’ Quarry specimens but from another Naracoorte
cave, Victoria Cave 5U1. Two scientists, who were also
avid cavers, were to make a remarkable discovery that
would highligh t the scientific impor ta nc e of the
Naracoorte Caves to the world.
Victoria Fossil Cave
Discovered by Reddan in 1894 and opened to the public
in 1897 (Anonymous 1897), Victoria Cave (now Victoria
Fossil Cave) is one of the main Naracoorte tourist caves.
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Top. Figure 3 - Specimens collected by Reddan and Zietz
in the South Australian Museum; Photo: Liz Reed.
Below. Figure 4 - Newspaper article recording the
Museum visit to Naracoorte Caves to collect Thylacoleo
specimens. The Register, Saturday December 12th
1908 pp. 4.
Page 9 ACKMA Journal No. 97 December 2014
During explorations in 1969, Grant Gartrell and Rod
Wells (members of CEGSA) pushed through a 10 m long,
low rocky passage located about halfway through the
known cave and found a chamber filled with countless
fossils. Limb bones, skulls and other elements could be
seen lying on the surface and poking through the
sediment (Wells, 1975). As Gartrell explored the cavern,
Wells scanned the sediment from a ledge at the front of
the deposit where he found an encrusted Thylacoleo
skull lying at his feet (Figure 6). Gartrell found another
skull lying on a rock pile about halfway along the
chamber (Wells, pers. comm. 2014). Over the next few
years, the remains of at least 18 individual Thylacoleo
were collected from the deposit (Wells et al., 1984). The
chamber contains a sediment cone, which was deposited
through an entrance hole located above the northeast
end of the chamber. That entrance is now blocked, but
once acted as a “pitfall” trap, collecting and preserving
large numbers of animals over a period of more than
200,000 years (Wells, 1984; Reed, 2008).
It would be difficult to overstate the significance of the
discoveries in Victoria Fossil Cave. The Main Fossil
Chamber is massive, some 54 metres long, about 16
metres at its widest point and at least 3 metres deep
(Reed, 2008). The accumulation dates from around
213,000 to as far back as 478,000 years ago (Reed and
Bourne, 2009). The sediment deposit contains a diverse
assemblage of fauna, with over 100 species of vertebrates
recorded (Reed and Bourne, 2000). Numerous important
Thylacoleo specimens have been recovered from this rich
deposit, including the articulated right and left manus
and an almost complete pes; which were recovered by
Wells soon after the chamber was discovered (Wells,
1975). Grant Hall, a chamber located just beyond the
Fossil Chamber and discovered in 1975, covers a period
from around 70,000 to 93,000 years ago (OSL
chronology), providing an almost complete chronology of
the late Pleistocene fauna (Macken et al., 2011). The
Upper and Lower Ossuaries, located through low
passages beyond the main Fossil Chamber, contain
breathtaking fossils littering the sandy-clay floor of a
chamber covering an area of around 450 m2 (Reed,
2006). The Ossuaries are so remarkable, that David
Attenborough filmed the cave for his “Life on Earth”
series, which aired in 1979 (Nolan, 1977, Figure 7).
He also mentioned them in his book of the same name
(Attenborough, 1979). As the chambers contain relatively
pristine sediment floors, they are protected as a
“Reference Area” for the Naracoorte Cave system – no
excavation has taken place in the chamber and no
research is allowed without special permission and strict
conditions. Fossils of Thylacoleo, Megalibgwilia,
Zygomaturus and Thylacinus are spread across the
surface, but they are overwhelmed by the number of
Sthenurine kangaroos – which account for almost 60% of
the visible material (Reed, 2006). Butch and Lake
Chamber, a small chamber adjacent to the Main Fossil
Chamber has also provided Thylacoleo fossil material.
An underground museum
The scientific value of the Victoria Fossil Cave deposits
was recognised at once by Rod Wells, who wrote in a
special report to the Deputy Director of the South
Australian Government Tourist Bureau strongly
recommending the area be protected, as “…it could yield
a considerable amount of scientific information if studied
by qualified zoologists and palaeontologists”. Furthermore
if properly developed it could also be of educational value
to the general public” (report by Wells September 1969 –
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Figure 6 - Thylacoleo skull in situ, Fossil Chamber,
Naracoorte Caves
Photo: Rod Wells
Figure 5 - James Quarry Thylacoleo manus specimen.
Photo of image held in the South Australian Museum
collection - Photo: Liz Reed
Page 10 ACKMA Journal No. 97 December 2014
R. Wells pers. comm. 2014). Tours in Victoria Cave had
occurred since its discovery and had focussed on the
impressive speleothem formations in the cave; but the
idea of the deposit being an underground museum for
communicating the scientific value of the fossil deposit
was novel, and one that would shape the future of
tourism at Naracoorte. In 1970, Dr Richard Tedford,
Curator of Vertebrate Palaeontology from the American
Museum of Natural History in New York visited the site.
He was quoted in the Canberra Times (July 18 1970
page 13) to have remarked - “From these exceptionally
rich deposits we may get some answer to the puzzle of
why so many forms of large animal life became extinct”.
Tedford also noted that “.. similar bone caves in the
United States and Europe had provided excellent tourist
attractions”.
By the mid to late 1970s, the Tourist Bureau had
installed electric lighting at the excavation area and was
conducting tours to the site for the general public. Part
of the ticket office at the caves was made available for
Rod and his team to use as a field laboratory and by
1971 Rod was working together with Ern Maddock of the
Tourist Bureau to develop displays for visitors. One of
these was a silhouette image of Thylacoleo, that Rod had
reconstructed based on the two known skeletons of the
time. The focus of guided tours shifted to discussions of
the significance of the deposit and on vertebrate
palaeontology of the late Pleistocene in Australia. A
workbench had been set up at the dig and fossils were
used as props by cave guides. The Tourist Bureau
improved facilities at the park in preparation for the
increased numbers of visitors the new site had attracted.
Currently a 60 minute tour runs at least twice daily
through Victoria Fossil Cave. The tour includes a 30
minute talk held in the Fossil Chamber, just above the
Excavation Site A (Wells et al., 1984), where a purpose
built viewing and seating area supports up to 25 visitors.
Immediately below the viewing platform, hundreds of
exposed skeletal elements are visible in the pit, relieved
out in situ. A full skeletal cast of Thylacoleo carnifex, as
well as Simosthenurus occidentalis, is mounted on
display at the front of the platform (Figure 8). The cast of
Thylacoleo is a composite model, reconstructed from a
number of individual specimens which have been
excavated from the Fossil Chamber. The hands and
forelimbs, pelvis and hind limbs of the model were cast
from those used by Wells and Nichol (1977). The
vertebrae were cast from individual bones found
associated with a Thylacoleo skull in the Fossil Chamber
(R. Wells, pers. comm. 2014). Casting of the manus and
pes was done by Rod Wells, limbs and vertebral column
by Ed Baily and the skull and dentaries by Peter Daenke
at Flinders University. The cast is posed in a somewhat
aggressive stance, and is striking and photogenic,
appearing in numerous photos on popular photography
websites, such as Flickr and is currently featured on the
Wikipedia entry for the Naracoorte Caves (2014).
Thylacoleo has also featured on various tourist websites,
brochures and information boards for the region (Figure
9a,b).
A second cast from the same material is on display at the
caves visitor centre, the Wonambi Fossil Centre, where
the reconstruction is “struggling” with the extinct
madtsoiid snake, Wonambi naracoortensis. This
reconstruction is the centre piece of the visitor centre
foyer, and accompanies two other reconstructions of
Thylacoleo; both animatronic, life models. The original
robotic model is located in the diorama, along with a
number of other life-sized Pleistocene reconstructions,
such as Diprotodon optatum, Zygomaturus trilobus and
Palorchestes azael. Artist Stephen Hayter made the
robotic models in consultation with Rod Wells and they
were constructed at Flinders University. The models were
revealed to the public when the centre opened in 1998.
The robot moves its head from side to side and growls.
The second model, located in the foyer near the mounted
cast, is a more recent addition, again made by Stephen
Hayter and installed around 2005.
The cast and life models are popular with tourists, but
lack the visual impact of the cast mounted in the Fossil
Chamber. The location, pose and atmosphere of the
reconstruction at the Fossil Chamber provide a depth
and dimension to the science of palaeontology that it is
not possible to recreate in a standard museum. The
guide’s talk, in front of the fossil bed, dwarfed by the
shadow of the Thylacoleo skeleton on the cave walls,
creates a powerful impression. The skeletons at the fossil
bed also provide important props to assist the guides in
conveying the process of palaeontology and how
scientists work from bones to reconstruct extinct
animals.
In 1994, Naracoorte Caves, along with the Riversleigh
Fossil Sites in North West Queensland, received
international recognition and protection under the
UNESCO World Heritage program. UNESCO ranks the
fossil sites as being “among the world's 10 greatest fossil
sites” and singles out Victoria Fossil Cave (5U1)
Naracoorte as being “… in terms of both volume and
diversity, Australia's largest and best preserved and one
of the richest deposits in the world” (UNESCO 2014).
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Figure 7 - David Attenborough with Thylacoleo skull in
the Ossuaries, Victoria Fossil Cave, Naracoorte
Photo: Rod Wells
Page 11 ACKMA Journal No. 97 December 2014
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Above. Figure 8 - Thylacoleo and a sthenurine kangaroo skeleton cast shadows on the wall in Victoria Fossil Cave
Below left. Figure 9a - Guide interacting with visitors using Thylacoleo skeleton. Cover of regional visitor guide
Photo: Steve Bourne; Below right. Figure 9b - 2005 park brochure featuring a Thylacoleo skull. Photo: Steve Bourne
Page 12 ACKMA Journal No. 97 December 2014
The fossil record of Thylacoleo at Naracoorte Caves
Victoria Fossil Cave is spectacular not just for the
number and size of fossil deposits in its numerous
chambers, but also for the excellent preservation of the
specimens that have been recovered. The left and right
manus of Thylacoleo, along with some of the forelimb
bones, were recovered fully articulated and preserved in
calcite (R. Wells pers. comm. 2014). These specimens
finally provided the reference material needed to fully
describe these aspects of the animal (Wells and Nichol,
1977). Analysis of the articulated manus showed
Thylacoleo was capable of a wide range of motion of the
first phalanx (tipped with a large recurved claw),
including gripping by opposing the pisiform. The other
phalanges were each tipped with ungual crests,
homologous to those found in felids, which allow the
animal to retract its claws. Thylacoleo also had a wide
range of motion of its forearms, and was capable of using
its hands for powerful grasping. Unfortunately, the pes
was incomplete, preventing a deep understanding of
Thylacoleo’s locomotion and stance.
In 1969, at a quarry owned by the Henschke family, a
cave was opened up during limestone quarrying. The
cave, later named Henschke’s Fossil Cave (5U91, 5U97)
was a substantial cave, first surveyed at about 50m long
with later extensions discovered adding an additional
60m of passages at two levels. Henschke Industries
kindly changed their quarrying plans and left the cave in
an accessible state until 1981, allowing palaeontologists
from the SAM to excavate and document the fossils.
Neville Pledge (Curator of Fossils), calculated there were
at least 46 individuals of Thylacoleo from the deposit,
making it more far more common that other carnivores
such as Thylacinus cynocephalus (12 individuals) and
Sarcophilus (6), and similar to the number of Sthenurine
kangaroos (52) (Pledge, 1990). The large mammal
component of the deposit was dominated by
macropodine kangaroos; with Macropus rufogriseus and
Macropus giganteus comprising 482 individuals. Pledge
noted that most of the Thylacoleo material recovered
from the cave system was from juveniles, mostly very
young individuals with un-erupted, or recently erupted
teeth. He attributed this to the pitfall nature of the cave
entrance and the inexperience of younger animals,
presumably hunting around the vicinity. The faunal
composition of the deposit was similar to that of Victoria
Fossil Cave; however, the stratigraphy and taphonomic
properties of the Henschke’s deposit were unclear. Given
the salvage nature of the excavation; the site did not lend
itself to a more thorough interpretation.
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Site
Description
Reference
Victoria Fossil Cave 5U1
Main Fossil Chamber
Excavation by R. Wells, E. Reed and others
Wells et al. (1984); Reed and Bourne
(2000, 2009).
Grant Hall
Excavation by R. Fraser and later A. Macken.
Fraser and Wells (2006); Macken et al.
(2012); Reed and Bourne (2000, 2009).
Upper and Lower
Ossuaries
Material collected from cave floor and
documented in situ.
Reed (2006); Reed and Bourne (2000).
Butch and Lake Chamber
Material collected from cave floor and
documented in situ.
Reed and Bourne (2000).
Bat Cave 5U2
Material collected from the entrance chamber
Tideman (1967); Maddock (1971); Reed
and Bourne (2009).
Alexandra Cave 5U3
Material collected by caves caretaker W.
Reddan.
Pledge (1977); Reed and Bourne
(2000).
Blanche Cave 5U4,5,6
Incidental finds in cave tunnels; later formal
excavation by T. Laslett and expanded by E.
Reed.
Reed and Bourne (2000, 2009, 2013).
Stick-Tomato Cave (Wet
Cave) 5U10, 11
Excavation in the entrance chamber of the
cave.
Reed and Bourne (2000, 2009).
Cathedral Cave 5U12, 13
Incidental finds in cave tunnels; subsequent
organized excavation in fossil chamber by S.
Brown and G. Prideaux.
Daily (1960); Brown and Wells (2000);
Prideaux et al. (2007).
Robertson Cave 5U17,
18, 19
Excavation in inner chamber of cave by M.
McDowell and later S. Brown.
Reed & Bourne (2009).
Table 1 – Fossil sites that have yielded remains of Thylacoleo carnifex - Naracoorte Caves World Heritage Area.
Page 13 ACKMA Journal No. 97 December 2014
In just a few years since these major discoveries,
Naracoorte had provided enough Thylacoleo material to
address some long standing taxonomic questions. Using
material contributed from Naracoorte and Wellington
Caves, Archer and Dawson (1982) began to revise the
taxonomy of the Thylacoleonidae. They found that skulls
va rie d gre atl y be twe en in div idu als i n th e sam e
geographic region and previous taxonomic distinctions
actually fell within the normal range. Owen’s
Thylacopardus australis was also found to fall within the
normal size range of Thylacoleo carnifex, as was Krefft’s
Thylacoleo robustus (Krefft 1872) and McCoy’s Thylacoleo
oweni (McCoy 1876). Krefft’s Mylodon australis ungual
(Krefft 1870) was also found to belong to Thylacoleo, as
was Krefft’s proposed genus Plectodon (Krefft 1870). The
controversial nature of the taxonomy of the genus
Thylacoleo had been neatly resolved to just one
Pleistocene species, Thylacoleo carnifex; and the question
of Thylacoleo’s dietary and locomotory habits could now
be addressed. Important biomechanical investigations of
the post-cranial skeleton were made during the 1980s
(Finch and Freedman, 1986 and 1988), setting the stage
for later discoveries at Naracoorte and elsewhere. Wroe et
al. (1999) used material from Naracoorte and elsewhere
to determine body weight of Thylacoleo.
Wells et al. (1982) used material from Victoria Fossil
Cave to resolve the century old question of Thylacoleo’s
dietary habits. Functional analysis of the jaw and tooth
dynamics of Thylacoleo showed the animal’s dentition
was capable of exhibiting forces and shearing required to
process large prey. Modelling of feeding behaviour also
showed that micro-wear patterns on the teeth matched
those generated by feeding on meat, rather than an
herbivorous diet. Further supporting evidence came from
another Naracoorte specimen from the Henschke’s
Quarry, where chemical analysis using Strontium (Sr)
and Zinc (Zn) ratios was used to show that Thylacoleo
was carnivorous (Nedin, 1991). The results of these
studies have not been challenged and more recent work
using finite element analysis techniques (Wroe et al.,
2005, 2008) have supported this finding, revealing that
comparatively, Thylacoleo had a higher bite force than its
eutherian namesake. Multivariate analysis of both
cranial and post cranial features of Thylacoleo show that
it clusters closely with other Pleistocene felid predators,
such as Smilodon (Wroe, 2008). Owen’s original
hypothesis, that Thylacoleo was a well-adapted carnivore
despite its ancestors being of a more herbivorous diet
and “of good conscience”, is now accepted theory.
In 2005 another cave was unearthed in the Henschke’s
quarry system, this time with a front-end loader, which
gave the new cave its name. Komatsu Cave (U240) was
opened around 11 metres below surface level and
contained sediment cones with numerous fossils; again
the Henschke family offered to change their plans to
allow palaeontologists to collect and document the
material. Two of the authors (LR and SB), assisted by
Friends of Naracoorte Caves volunteers and students
from Flinders University, conducted a salvage operation
to excavate the site. The cave contained the remains of
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Site
Description
Reference
James’ Quarry Cave 5U29
Small cave uncovered during quarrying; fossils collected
by N. Tindale and P. Lawson (SAM).
Daily (1960); Pledge (1977);
Reed and Bourne (2000).
Henchke’s Fossil Cave 5U91,
5U97
Cave uncovered during quarry activity. Excavated by
SAM palaeontologists and later J. Barrie.
Pledge (1990); Barrie (1997);
Reed and Bourne (2000).
Komatsu cave 5U240
Cave uncovered during quarry activity. Excavated by S.
Bourne, E. Reed and others.
Reed and Bourne (2009).
Komatsu Cave 2
Cave uncovered during quarry activity. Excavated by S.
Bourne, E. Reed and others.
Reed and Bourne (2009).
S102 Cave 5U47
Isolated pes elements found in cave tunnel by S. Bourne
Reed and Bourne (2009).
Haystall Cave 5U23
Fossil material collected from the sediment of a large
cone.
Reed and Bourne (2000).
Specimen Cave 5U35 (also
known as Zietz Cave)
Fossil material collected from sediments and cave
tunnels by W. Reddan and E. Stirling.
Stirling (1908, 1912); Reed
and Bourne (2000).
Cable Cave 5U125
Fossil material collected from cave floor.
Reed and Bourne (2000).
Buckridge Cave 5U169
Small cave uncovered and subsequently destroyed
during vineyard preparation. Excavated by S. Bourne,
E. Reed and M. McDowell.
Reed and Bourne (2000).
Crawford’s Cornucopia Cave
5U171
Small cave uncovered during vineyard preparation.
Excavated by S. Bourne and E. Reed
Reed and Bourne (2000).
Whale Bone Cave 5U250
Cave uncovered during vineyard preparation; sediment
floored chamber with fossil material evident.
Reed and Bourne (2009).
Table 2 – Cave sites within the Naracoorte District* that have yielded Thylacoleo material.
*Other sites with Thylacoleo carnifex in South East Region - Comaum Forest Cave 5U118, Green Waterhole Cave
5L181, Wandilo Forest Cave 5L365, Gouldens Hole 5L8, Un-named Cave 5L441, Moorak, Kilsby’s Hole 5L46, Un-
named cave in Mount Gambier (site 45 in Reed & Bourne 2000).
Page 14 ACKMA Journal No. 97 December 2014
both juvenile and adult Thylacoleo (Figure 10). For the
first time, an articulated pes (complete with hind limb
attached), was recovered (Figure 11). Using this fossil,
Wel ls et al. (2009) were able to reconstruct the
morphology of the pes, completing the story of
Thylacoleo’s limb morphology. Thylacoleo had
morphological elements of both a scansorial quadruped,
and adaptations consistent with a climber.
Based on their interpretation, Thylacoleo would have
been able to grasp branches with its foot, and thrust its
body weight upward in a manner of climbing similar to
modern possums (Wells et al., 2009). Thylacoleo has
been found across Pleistocene fossil cave deposits in
both the Naracoorte World Heritage Area (see Table 1)
and the greater Naracoorte district (Table 2), spanning a
time period of at least 300,000 years. It has now been
found across fossil deposits in all states of Australia, and
in many bioregions and environments (Prideaux, 2006).
The regularity of its inclusion in these deposits is
interesting, as large carnivores are usually
underrepresented in fossil deposits when compared with
other animals such as kangaroos. Thylacoleo may have
been attracted to the caves to feed on trapped or injured
animals, creating a similar accumulation bias to that
seen in deposits such as the Le Brea Tar Pits in Northern
America (eg. Spencer and Van Valkenburgh, 2003).
Megafauna icon
Larg e, c arn ivoro us a ni mals tend to captur e the
imagination and a ‘marsupial lion’ is unique and
intriguing to scientists and the general public alike.
Reconstructions of the animal have changed over time in
light of new scientific evidence. Many of these discoveries
have inspired popular articles, documentaries and
memorabilia. In 2008, Australia Post published a series
of stamps depicting Australian megafauna species. The
stamps were designed by Peter Trusler, an accomplished
palaeoartist, and featured Thylacoleo carnifex on the 55c
stamp. In 2014, Perth mint released a 1oz silver coin
depicting an adult Thylacoleo with a young on its back.
This coin, part of a limited release, was also designed by
Trusler. In 2011, for the Perth Mint coin release
“Australian Fossil Mammal Sites” “World Heritage Area”,
Thylacoleo was chosen to represent Naracoorte Caves.
In recent years, Thylacoleo has been prominent in media
appearances and scientific interpretation at the
Naracoorte Caves. Many of these are filmed at the fossil
pit under the shadow of Thylacoleo. In 2009, the cast
skeleton at the fossil bed was used to discuss the habits
of Thylacoleo for the Discovery Channel production
“Monsters Resurrected, Episode 6 – Megalania, Giant
Ripper” which aired in October 2009 and included one of
the authors (LR). This is just one example from many. A
photograph (taken by one of the authors SB) of a young
girl posing with the Thylacoleo skeleton at the fossil bed,
quickly became the ‘face’ of Geo-tourism in South
Australia and was used in multiple tourism promotions,
advertisements and tourism award nomination
documents (Figure 12).
In 2010, an article in National Geographic Magazine
(“Australia’s Lost Gi an ts ”) , inclu de d a lif e- like
reconstruction of Thylacoleo carnifex created by the
Kennis brothers. This was based on a skull from
Komatsu Cave in Henschke’s Quarry at Naracoorte.
Many visiting palaeontologists have been photographed
with the Thylacoleo skeleton at the fossil bed, including
Dr Phil Currie and Dr Jackson Njau (Figure 13).
The End of the Tail
Naracoorte has played a pivotal role in both the scientific
understanding and public profile of the extinct marsupial
lion. Thylacoleo fossils have now been recovered from
every state in Australia and from various biogeographic
RESEARCH
Figure 11 - A reconstructed hind limb and pes of
Thylacoleo from Komatsu Cave, Naracoorte.
Photo: Steve Bourne
Figure 10 - Cath Sellars excavating a Thylacoleo skull in
Komatsu Cave 2, Naracoorte. Photo: Steve Bourne
Page 15 ACKMA Journal No. 97 December 2014
regions and climates. In 2002 on the Nullarbor Plain, a
team of cavers from the Victorian Speleological
Association (VSA) were systematically surveying the
Nullarbor Plain in Western Australia when they
discovered a number of previously unknown cave
entrances. During surveying of the new caves, they
found the almost complete, articulated skeleton of
Thylacoleo, exquisitely preserved, lying on the sediment
in a chamber as if it had recently expired. The find was
reported to the Western Australia Museum (WAM) and
collected soon after by palaeontologists including Gavin
Prideaux and John Long from Flinders University. The
skeleton provided something that had eluded
palaeontologists for 150 years – a tail.
Tails play an important part in the locomotion of
marsupials; not just in climbers such as possums.
Macropods use the tail as a ‘fifth leg’ in pentapedal
locomotion. Tasmanian devils and other dasyurids use
the tail to facilitate a ‘tripod’ stance, allowing these
carnivores to stand on their hind legs. Current work by
P r o f ess o r R od We l ls a n d c ol lea g u e s in v o l ve s
biomechanical analyses of the vertebral column of
Thylacoleo skeletons from the Nullarbor Caves and
Naracoorte. This will allow insight into the gait and
stance of this enigmatic marsupial, putting in place the
final pieces of the Thylacoleo puzzle.
Acknowledgements
The authors thank Professor Rod Wells for allowing us to
use some of his photographs and for personal accounts
of discoveries and interpretations of fossil material at
Naracoorte Caves.
RESEARCH
Figure 12 - Young girl posing with the Thylacoleo
skeleton in Victoria Fossil Cave. Photo: Steve Bourne
Figure 13 - Dr Phil Currie (above) and Dr Jackson Njau
(below). Photo: Steve Bourne
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