SUAS tytn BTN or iy i ate Por wie ght , Spa ‘ hp etree 5 y : wee Uf read hay e ie Bye the naire rans ; $ eed r SuENaNaNf a8 dM uw it Peter regs soe fey Laatste Sai Fathi sh tute an he ENS Jrathte twk Rice ial SLES 2 , ; ayo) oni “ay Fake dere i ni Sala ; rod eh Jejnets fe By ae ; RAN Tar ycitytte bet . eth ot ae ye ‘ thorn de ae nthe ge Pater d eer Laie, PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES This volume contains a selection of papers dealing with vertebrates or vertebrate fossil sites, arising from a Symposium held by the Linnean Society of New South Wales at Wellington Caves, New South Wales, in December 1995. Papers concerning Holocene vegetation will appear in volume 118. The opening session of that symposium was held in Cathedral Cave. The cover of this volume is based on a colour photograph, supplied by the Museum of Natural History, London, of an aquatint of nearby Breccia Cave, also known as Mitchell’s Cave, by Thomas Mitchell. It was first published in 1831 in Part XIV of the Edinburgh New Philosophical Journal. A version of this drawing was also published by Mitchell in 1838 in his book “Three expeditions into the Interior of Eastern Australia”. The legend to the original states, “The cave in which the fossil bones are found at Wellington Valley is in compact secondary limestone, as described more fully in a memo- randum which accompanied a very large bone sent by Mr Rankin to Prof. Jameson. It is near a larger cave* where no breccia has been found, and which is very different in character from that of which this drawing is intended to convey an idea, the appearance of disruption and with unshapely masses overhanging being characteristic of all the situations where the fossil bones have hitherto been discovered in N.S. Wales. 12 Oct 1830. The bone above mentioned is that of an elephant**”’. *Cathedral Cave **Tt was Diprotodon VOLUME 117 March 1997 Dedication PROFESSOR RICHARD DEHM July 1907— March 1996 The contents of this volume would have been noticeably different and the course of research into Australian Quaternary mammals substantially altered had fate not inter- vened to prevent research begun by Dr Richard Dehm from being concluded. Inspired by lectures and contacts with cave researchers from his teenage years, Dr Dehm jumped at the chance to travel to Australia in 1939 with his colleague Dr Joachim Schréder, sup- ported and encouraged by the director of the Bavarian Collection of Palaeontology and Historical Geology in Munich. The timing was however most unfortunate, and the two scientists found themselves in Australia when war broke out between Germany and Great Britain. The story of their internment and difficult return to Germany has been partly told (Augee at al., 1986, The Australian Zoologist 22, 3-6), and further details and documentation provided by Prof Dehm shortly before his death will, when fully translat- ed, be incorporated into a future publication. The tragic outcome for palaeontology was that fossils collected from Australia, especially from Wellington Caves and Jenolan, were largely destroyed or lost. The few that survived the misfortunes of war remain in the museum at Munich, excellently preserved and stored, but are of little research value. A list which I compiled while on study leave in Munich in 1986 is held in the archives at the Wellington Caves Fossil Studies Center. Richard Dehm had the view that the remains of small animals, particularly mam- mals, held the secret to understanding climatic and faunal changes. He studied fossils found in fissure fillings, and therefore Wellington Caves were of particular interest. Prof Dehm had a life-long desire to return to Australia and sites such as Wellington and Jenolan, however a successful career and the added duties from 1958 as Director of the “Universitatsinstitut and Staatssammlung fiir Palaontologie und Historische Geologie Miinchen” intervened and he was never to make the journey. In a way the research he would have carried out was finally begun in the early 80s when the University of NSW began the stratigraphically controlled excavation in Cathedral Cave. A brass plaque at Wellington Caves marks one of the spots from which the remaining Munich collection was obtained, and this is readily viewed by visitors to the Phosphate Mine section. Prof Dehm remained very active following his retirement as Director of the Munich collection and museum. During my three month study of the Australian mammal fossil in Munich, he was a delightful host, with a keen interest in current Australian palaeontological studies on which he was well informed. His death marked the end of a remarkable career and a sad loss to colleagues around the world. M.L. Augee Richard Dehm (left) and Michael Augee on the steps of the Munich museum which houses the Bavarian State fossil collection, winter 1985. Proc. LINN. Soc. N.S.w., 117. 1997 z a es 9 oer Huis stati y Say Ciel tbo) bs reqate oh Wey Wogan 1S mbyorciee piel A sy wives Ws ental, or gaseeiaillaraett Giant Ringtail Possums (Marsupialia, Pseudocheiridae) and Giant Koalas (Phascolarctidae) from the Late Cainozoic of Australia MICHAEL ARCHER, KAREN BLACK AND KERRY NETTLE School of Biological Science, University of New South Wales, Sydney N.S.W. 2052 Archer, M., Black, K. and Nettle, K. (1997). Giant Ringtail Possums (Marsupialia, Pseudocheiridae) and Giant Koalas (Phascolarctidae) from the late Cainozoic of Australia. Proceedings of the Linnean Society of New South Wales 117: 3-16 While seven pseudocheirids have been described from the late Oligocene to early Miocene and five from the Pliocene of Australia, none are known to have been confined to the Pleistocene. We describe here a giant ringtail, Pseudokoala cathysantamaria n. sp., from the early Pleistocene Portland Local Fauna (lower Nelson Bay Fm) of Victoria. We review the generic distinction of the Miocene—Pliocene Corracheirus Pledge, 1992 and conclude that it is a junior synonym of Pseudokoala Turnbull and Lundelius, 1970. The palaeohabitat for all species of Pseudokoala appears to have been rainforest. A second pseudocheirid in the Portland Local Fauna is most similar in morphology and size to the still-living Pseudocheirus peregrinus, a species whose habitat range also includes rainforest. We describe the first known upper and additional lower molars for the giant koala Cundokoala yorkensis Pledge, 1992, from a Pleistocene deposit in Wellington Caves, New South Wales, a taxon previously only known from Pliocene sediments of South Australia. The Portland giant ringtail and Wellington giant koala add two more families to the already extensive list that exhibited Pleistocene gigantism. Previously, only one other lineage (tree kangaroos) of arboreal mam- mals has been known to have undergone gigantism. Manuscript received 6 June 1996, accepted for publication 23 October 1996. KEYWORDS: Koalas, Phascolarctidae, Pleistocene gigantism, Pseudocheiridae, Quaternary, ringtail possums. INTRODUCTION The record for ringtail possums (Pseudocheiridae) begins in the late Oligocene. From sediments of this age in South Australia, Woodburne et al. (1987a) and Pledge (1987b) describe four species of Pildra, two of Marlu and one of Paljara. Archer (1992) reports over twelve additional unnamed taxa from the Oligo-Miocene sediments of Riversleigh, northwestern Queensland. Archer and Bartholomai (1978) note a Pseudochirops from the late Miocene of Alcoota, Northern Territory. Turnbull and Lundelius (1970; augmented by Turnbull et al. 1987) describe Pseudokoala from the early Pliocene Hamilton Local Fauna as well as two species of ‘Pseudocheirus’ which may (Archer 1984) represent the otherwise modern genus Petauroides. Another species of Pseudochirops, from the early Pliocene Bluff Downs Local Fauna of northeastern Queensland, is under study (B. Mackness and M. Archer). Pledge (1992) describes a giant species of Corracheirus from the late Miocene to Pliocene Curramulka Local Fauna of southern South Australia as well as a single tooth of another large pseudocheirid from the Plio-Pleistocene Cement Mills Local Fauna of southeastern Queensland. Until now, how- ever, no giant pseudocheirids have been reported from undoubted Pleistocene sediments. The tooth described here as Pseudokoala cathysantamaria, from early Pleistocene Proc. LINN. Soc. N.S.W., 117. 1997 4 GIANT RINGTAIL POSSUMS AND GIANT KOALAS deposits at Portland, Victoria, is the only ringtail possum confined to the Pleistocene and the only giant ringtail known from the Quaternary. Fossil koalas (Phascolarctidae) are similarly known from late Oligocene and younger sediments. Stirton (1957), Stirton, Tedford and Woodburne (1967), Woodburne et al. (1987b) and Springer (1987) describe six species in the Oligo-Miocene genera Perikoala, Litokoala and Madakoala. Black and Archer (in press) name another Miocene genus from Riversleigh and Black (in prep.) reports additional Miocene taxa from Riversleigh. Pledge (1992) describes a Miocene to Pliocene species of the giant Cundokoala. Bartholomai (1968) describes a large Plio-Pleistocene Queensland species of the modern genus Phascolarctos. Pledge (1987a) describes another large Phascolarctos from the early Pliocene of South Australia. We describe here isolated teeth, from Pleistocene deposits of Wellington Caves, New South Wales, that appear to represent the same Miocene/Pliocene South Australian species described by Pledge (1992) as Cundokoala yorkensis. The Wellington material includes the first known upper molars for this taxon. Material described is registered in the palaeontological collections of three institu- tions as follows: AM F numbers represent the Australian Museum fossil collection; P numbers represent the palaeontological collections of the Museum of Victoria; SAM P numbers represent the palaeontological collections of the South Australian Museum. Dental terminology used here follows Luckett (1993) for molar homology, Flower (1867) for premolar homology and Archer (1984) and Woodburne et al. (1987a,b) for molar morphology. SYSTEMATICS Class Mammalia Linneaus, 1758 Superorder Marsupialia [liger, 1811 Order Diprotodontia Owen, 1866 Superfamily Phalangeroidea Weber, 1928 Family Pseudocheiridae Winge, 1893 Pseudokoala Turnbull and Lundelius, 1970 Type species: Pseudokoala erlita Turnbull and Lundelius, 1970 Additional species: Pseudokoala curramulkensis (Pledge, 1992) and Pseudokoala cathysantamaria n. sp. Revised generic distribution: Corra Lynn Cave, Yorke Peninsula, South Australia; Hamilton Local Fauna, south- western Victoria; Portland Local Fauna, Nelson Bay Formation, Portland, Victoria. Revised age range for the genus: The Hamilton Local Fauna is interpreted (Rich et al. 1991) to be early Pliocene in age. The Curramulka Local Fauna is interpreted (Pledge 1992) to be late Miocene to Pliocene in age; the Nelson Bay Formation is interpreted (Flannery and Hann 1984) to be early Pleistocene in age. Revised generic diagnosis: Species of Pseudokoala are distinguished from Marlu praecursor by their continu- ous postmetacristid and preentocristid, and posthypocristid which does not extend to the Proc. LINN. SOC. N.S.W., 117. 1997 M. ARCHER, K. BLACK AND K. NETTLE 5 lingual side of the crown. They are distinguished from M. kutjamarpensis by their con- tinuous postmetacristid and preentocristid, connection of the cristid obliqua to the metacristid and their truncated posthypocristid. They are distinguished from species of Pildra and Paljara in their lack of an entostylid, presence of a buccal buttress on the pro- toconid, en echelon overlap of the postmetacristid and preentocristid, large size, and truncated posthypocristid. They are distinguished from species of Pseudocheirus, Petauroides and Hemibelideus in having a truncated posthypocristid, being much larger in size, having a second buccal buttress on the protoconid, not having steep entoconid blades and in lacking a protostylid. They are distinguished from species of Pseudochirops in having a truncated posthypocristid, being larger in size, having a sec- ond buccal buttress on the protoconid, not having steep entoconid blades, having the cristid obliqua connected to the metacristid, and in lacking a protostylid. Pseudokoala curramulkensis (Pledge, 1992) Emendation of dental homology for the paratype and referred specimen: Pledge (1992) notes three specimens including the Holotype. The homology of the teeth (using the M1-4 molar homology system of Luckett 1993) given in table 5 needs emendation as follows. SAM P29901 (referred specimen) is a right dentary fragment with M> (not Mj) and alveoli for M; and M3_4. SAM P31792 (Paratype) is a left den- tary fragment with M>_3 (not Mj_>) and alveoli for M, and Mg. Pseudokoala cathysantamaria n. sp. Holotype: P173650, an isolated LM, (Fig. 1). Type Locality: Interpreted to be Nelson Bay, Portland, Victoria, although no specific locality details are recorded in the Museum of Victoria Palaeontology register (B. Thompson, pers. comm. to M. Archer), nor are there any details on the specimen label. However, it is regis- tered within a series of fossils, including P173649 (a dentary of Pseudocheirus sp. cf. P. peregrinus; see below), for all of which the locality data is Nelson Bay, Portland, Victoria. Age: Marine invertebrates from the Lower Nelson Bay Formation, the probable source of the Holotype, are early Pleistocene in age (Flannery and Hann 1984). Etymology: In honour of Cathy Santamaria for her constant interest and much appreciated encouragement for palaeontological research in Australia. TABLE | Measurements of Pseudokoala material. All measurements in millimetres. Specimen P3 MI M2 M3 M4 EW, a AWE Wan Iad AWE
_4, and SAM P29901 with M> which support the distinction with the
postprotocristid appearing to make no contact with the metastylid; and SAM P31792
with M>_3 where the postprotocristid bypasses (but is touched by) the cristid obliqua to
make contact with the metastylid flexure. Consequently, this feature appears to be vari-
able and does not distinguish species of Corracheirus from those of Pseudokoala.
Considering the third distinguishing feature, the Holotype and SAM P29901 do support
the suggestion that the preentocristid is continuous with metacristid via a simple
metastylid flexure. However, the condition in SAM P31792 is much closer to the P. erlita
condition in that there is only a very tenuous direct connection linking the preentocristid
and the metacristid via a fine and low bridge of enamel buccal to the metastylid. In this
specimen, the preentocristid and metacristid really overlap en echelon, with the preen-
tocristid passing anterobuccal to the posterior end of the metacristid to make contact with
the postprotocristid, a condition closer to that seen in P. erlita.
Proc. LINN. Soc. N.S.W., 117. 1997
GIANT RINGTAIL POSSUMS AND GIANT KOALAS
. LINN. SOC. N.S.W., 117. 1997
M. ARCHER, K. BLACK AND K. NETTLE 9
Conversely, comparison of SAM P31792 with the M3 or g of P. erlita illustrated in
Turnbull, Rich and Lundelius (1987, Fig. 2C) suggests basic overall similarity in all key
features (except, possibly, the degree to which the metacristid is lingually concave and
the degree of posterior development of the postentocristid; see below). Corracheirus cur-
ramulkensis, P. erlita and P. cathysantamaria also exhibit a striking synapomorphy of
Mj: truncation of the posthypocristid such that it stops well short of the lingual side of
the crown and does not contact any other structure. This condition differs from all other
pseudocheirids where the posthypocristid closely approximates the postentocristid (or the
entostylid) at or near the lingual margin of the tooth. All three taxa also exhibit a tenden-
cy to posteriorly extend the postentocristid, a feature best-developed in M}. Finally, all
three taxa appear to be united in their tendency towards gigantism. For these reasons, we
suggest that Corracheirus Pledge, 1992 is better regarded as junior synonym of
Pseudokoala Turnbull and Lundelius, 1970 which then contains three species: erlita
Turnbull and Lundelius, 1970; curramulkensis Pledge, 1992; and cathysantamaria
Archer, Black and Nettle (this paper), erlita being the type species of the genus.
Within Pseudokoala, P. cathysantamaria appears to be the sister-group of P. curra-
mulkensis, these two sharing as synapomorphies extreme gigantism and greater posterior
development of the postentocristid. Although the trigonid 1s unknown for P. curra-
mulkensis, that of P. cathysantamaria shows two prominent buccal buttresses in contrast
to one in P. erlita, the second buttress being an autapomorphic condition (or perhaps a
synapomorphic condition shared with P. curramulkensis). The single buttress of P. erlita
may be the homologue of the vertical protostylid ridge in the same position in species of
Marlu (or the homologue of the protostylid in, e.g., species of Pseudochirops,
Pseudocheirus, Pseudocheirulus).
Species of Pseudokoala share distinctive features of M, with species of Marlu
Woodburne, Tedford and Archer, 1987 including: failure of the cristid obliqua to directly
contact the postmetacristid (a deep crevice intervening); a crenulated cristid obliqua; en
echelon overlapping of the postmetacristid and the preentocristid (in M. kutjamarpensis
but not M. praecursor); a vertical buttress but no protostylid on the buccal flank of the
protoconid; no entostylid; and gently inclined (rather than steeply inclined) entoconid
blades. This intergeneric relationship was originally suggested by Woodburne, Tedford
and Archer (1987; Fig. 23) on the basis of synapomorphies exhibited primarily by the
upper molars. The new species P. cathysantamaria exhibits the same combination of
synapomorphies, lending support to the hypothesis of a Pseudokoala/Marlu clade.
Marked differences between species of Marlu and Pseudokoala include the trun-
cated posthypocristid of species of Pseudokoala and their better-developed entoconid
blades. Differences in the nature of the connection between the preentocristid and
metacristid vary within at least P. curramulkensis (see above) making intergeneric con-
trasts here of questionable value.
In terms of patristic relationships, the early Miocene Marlu kutjamarpensis
exhibits no features that would preclude it being ancestral to the Pliocene/Pleistocene
species of Pseudokoala.
Pseudocheirus Ogilby, 1837
Pseudocheirus sp. cf. P. peregrinus (Boddaert, 1785)
Specimen:
P173649, a right dentary with Mj_4 (Fig. 1).
Locality:
Nelson Bay, Portland, Victoria. Label data also records: ‘Monash Univ. field trip
(1973?) and “# 280 004’.
Proc. LINN. Soc. N.S.W., 117. 1997
10 GIANT RINGTAIL POSSUMS AND GIANT KOALAS
Age:
The marine invertebrate assemblage of the Lower Nelson Bay Formation, source
of P173649, is early Pleistocene in age (e.g., Flannery and Hann 1984).
Description:
P173649 is a Pseudocheirus peregrinus-sized ringtail possum (Table 2) with over-
all similarity to P. peregrinus and P. occidentalis. It is, however, distinguished from these
modern species (most individuals but not all, these features being somewhat variable in
at least P. occidentalis) in that: the preentocristid is connected to the metastylid on M>_4.
and the metaconid on My is less distinct. It is distinguished from Pseudocheirulus her-
bertensis in that: the preentocristid is connected to the metastylid; the M, lingual
parastylid is not developed; and there is no notch between the postmetacristid and
metastylid. It is distinguished from P. caroli and P. forbesi in that: the preentocristid is
connected to the metastylid; the Mj, lingual parastylid is not developed; the preen-
tocristid is strongly bladed; and there is no notch between postmetacristid and metastylid.
It is distinguished from P. canescens and P. mayeri in that: the preentocristid is connected
to the metastylid; the preentocristid is strongly bladed; and there is no notch between
postmetacristid and metastylid. It is distinguished from Hemibelideus lemuroides in that:
the buccal shelf is not prominent; and the M, metaconid is less distinct. It is distin-
guished from Petauroides volans in that: the preentocristid is connected to the
metastylid; there is no entostylid ridge; and the M; metaconid is less distinct. It is distin-
guished from Pseudochirops albertisii in: not having a protostylid basin on Mj; lacking
a prominent buccal shelf; lacking an entostylid ridge; lacking a posterobuccal trigonid
basin; and in having a less distinct My metaconid. It is distinguished from P. corinnae in
that: the preentocristid is connected to the metastylid; it lacks a protostylid basin on Mj;
it lacks a prominent buccal shelf; it lacks an entostylid ridge; it has no posterobuccal
trigonid basin; and the M, metaconid is less distinct. It is distinguished from P. cupreus
in: not having a protostylid basin on M}; lacking a prominent labial shelf, the My cristid
obliqua is not angulate, no entostylid ridge, no posterobuccal trigonid basin; and in hav-
ing a less distinct metaconid on M}. It is distinguished from P. archeri in: not having a
protostylid basin on Mj; having an Mg cristid obliqua that is not angulate; lacking an
entostylid ridge; lacking a posterobuccal trigonid basin; and in having a less distinct
metaconid on Mj}. It is distinguished from Petropseudes dahli in: not having a proto-
stylid basin on Mj; lacking a prominent buccal shelf; lacking a posterobuccal trigonid
basin; and in having a less distinct metaconid on Mj. It is distinguished from the species
of all extinct ringtail genera in having: a well-developed My protostylid; a preentocristid
connected to the metastylid; a strongly bladed preentocristid; an entoconid that is posi-
tioned anterior to the hypoconid; a protoconid positioned lingual to the midline of My;
and a less distinct metaconid on M}).
TABLE 2
Measurements of Pseudocheirus material. All measurements in millimetres.
Specimen P3 MI M2 M3 M4
LE W L AW PW L AW PW L AW PW L AW PW
MY P173649 AMG) SK ZA AV) AI) 230) AO), Dolls) D2 EB) AK} 21S)
AM M4046 3.51 1.58 Syke) IA, DP 3.84 2.12 2.54 4.03 2.37 2.60 4.74 2.43 2.30
The early Pleistocene habitat of Portland:
The presence of an ektopodontid and Pseudokoala cathysantamaria in the Portland
assemblage suggests a rainforest component in the palaeoenvironment. All ektopodon-
Proc. LINN. SOC. N.S.W., 117. 1997
M. ARCHER, K. BLACK AND K. NETTLE 1]
tids occur in assemblages that have been interpreted (e.g. Archer, Hand and Godthelp
1995) to represent rainforest communities or communities that include rainforest: the late
Oligocene Ditjimanka and Ngama Local Faunas, South Australia; the early and middle
Miocene assemblages of Riversleigh, Queensland; the ?early or middle Miocene
Kutjamarpu LF, South Australia; and the early Pliocene Hamilton LF, Victoria. The late
Miocene to Pliocene Curramulka LF of South Australia, which contains Pseudokoala
curramulkensis, is also regarded (Pledge 1992) to represent wet sclerophyll forest and/or
rainforest, as is the Hamilton assemblage (Turnbull and Lundelius 1970, Turnbull, Rich
and Lundelius 1987) which contains P. erlita. The occurrence in the Portland assemblage
of Pseudocheirus sp., cf: P. peregrinus does not conflict with this interpretation because,
although primarily an inhabitant of dense understorey vegetation, some modern popula-
tions of P. peregrinus extend well into rainforest.
Order Diprotodontia Owen, 1866
Suborder Vombatiformes Woodburne, 1984
Infraorder Phascolarctomorphia Aplin and Archer, 1987
Family Phascolarctidae Owen, 1839
Cundokoala yorkensis Pledge, 1992
Holotype:
SAM P24904, left dentary with Mj_4 and alveolus for P3.
Referred material
(Fig. 2) and (Table 3): SAM P24905, partial left Mz in its alveolus.
AM F98885, L MI: AM F98886, R M;; AM F98887, L Mj; AM F98888, L Mp;
AM F98889, R My; and AM F98890, L M3.
TABLE 3
Measurements of all known specimens of Cundokoala yorkensis. All measurements in millimetres.
Abbreviations: L, length; AW, anterior width; PW, posterior width.
Specimen p3 M! M2 M3 1
L WwW L AW PW L AW PW L AW PW L AW PW
AM F98885 12 I2Be 0
P3 M M> M3 My,
ee Wie Ihe AW PWat sees ar AWARE tenet MAW PWe te iMilnen AWA Wy
P24904 mS 720 8 “TO el TO Ra ey ae
AM F98886 mS 65 G0
AM F98887 11.5 6.9 68
AM F98888 MA 72 7.0
AM F98889 ms 7 GO
AM F98890 WAs Fos) @.7/
Revised distribution:
Cundokoala yorkensis is known from Corra Lynn Cave, the Curramulka Local
Fauna, Yorke Peninsula, South Australia and now Wellington Caves, New South Wales.
Proc. LINN. Soc. N.S.W., 117. 1997
12 GIANT RINGTAIL POSSUMS AND GIANT KOALAS
Revised age range:
The Curramulka Cave site of the typical material is interpreted by Pledge (1992) to
be Miocene to Pliocene in age. The exact locality and age for all but one (AM F98886)
of the Wellington Caves specimens is unknown. They were collected from the surface of
spoils piles dumped outside the newly restored entrance to the Phosphate Mine. The
material on these piles had been obtained during excavation of tourist paths in the
Wellington Caves complex in late 1995. The source of the material for each pile was not
certainly known but thought by excavation workers at the site to be various localities
within the Phosphate Mine and Bone Cave. Although most of the taxa recovered to date
from the Wellington Caves complex of sediments have been interpreted to be Pleistocene
in age, Hand, Dawson and Augee (1988), L. Dawson et al. (in prep.) and Osborne (1983)
have demonstrated that some deposits in the Wellington Caves complex (e.g., in the
entrance doline of Big Sink and others in the Phosphate Mine) are Pliocene in age.
Therefore the age of the Cundokoala yorkensis material obtained from the spoils piles
could be Pliocene, Pleistocene or both.
AM F98886, however, was obtained by one of us (MA) in company with H.
Godthelp, A. Gillespie, A. Musser et al. from a newly-excavated pathway in the Phosphate
Mine. Material excavated at the same time included an isolated lower molar of Diprotodon
sp. cf: D. optatum (only known from the Pleistocene), a dentary of Protemnodon sp. cf. P.
roechus (only known from the Pleistocene), abundant dentary and maxilla fragments of
Aepyprymnus rufescens (Pleistocene and living species) and a dentary of Onychogalea
unguifera (living species). Although other taxa obtained at the same time have yet to be
identified, nothing contradicts a Pleistocene age for this assemblage. We would conclude,
pending a thorough analysis of the rest of the fauna from this deposit, that Wellington Caves
C. yorkensis 1s Pleistocene in age, giving the species a Pliocene-Pleistocene age range.
Revised diagnosis
Cundokoala yorkensis differs from all other phascolarctids: in being larger; having
higher-crowned teeth; and in having a relatively short, massive dentary. It differs from all
other phascolarctids except species of Phascolarctos in having: a larger paraconule and
neometaconule on M°*; a more lingually positioned protoconid and a larger protostylid
on Mj; well-developed lingual columnar stylids on the metaconid and entoconid of Mq_
4; and a well-developed buccal cingulum and metastylid fold on Mj_y4. It differs from
species of Litokoala in: lacking the posterobuccal crest which extends from the apex of
the metaconid in Mj_ 4: lacking an anteriorly displaced entoconid (relative to the
hypoconid) on Mg; lacking an anteriorly displaced metaconid (relative to the protoconid)
on the My; and in having | the postprotocristid and cristid obliqua of My meeting in the
transverse median valley on My (in contrast to the parallel arrangement seen in Litokoala
kanunkaensis) It differs from species of Madakoala and Perikoala: in having: a larger
PUSH on M’; a more crenulate, less linearly-oriented paraconule and neometaconule
on M°; a paraconule that connects anterobuccally to the anterior cingulum on M°; an
entostylid ridge on Mj_4; a more lingual junction of the postprotocristid and cristid obli-
qua; and in lacking protoconid-metaconid and hypoconid-entoconid crests.
Description
The new Wellington Caves materials (AM F98885 to AM F98890) augment under-
standing about the morphology of this species. AM F98885 is an unworn, relatively square
selenodont M° that tapers posteriorly. It is morphologically very similar to the M* of
species of Phascolarctos and differs mainly in being relatively wider anteriorly. The para-
cone and metacone are similar in height and are the tallest cusps on the tooth followed by
the metaconule and protocone. The apex of the protocone lies lingually opposite that of the
paracone. The apex of the metaconule lies lingually opposite and slightly anterior to that of
the metacone. The lingual bases of the paracone and metacone and the anterolingual base
Proc. LINN. SOc. N.S.W., 117. 1997
M. ARCHER, K. BLACK AND K. NETTLE 13
of the protocone are highly crenulate. The protocone and metacone and their associated
crests are slightly obliquely (anterolingually) oriented. The tooth is bisected bucco-lingual-
ly by a deep transverse median valley and antero-posteriorly by a relatively deep longitudi-
nal valley. The buccal tooth margin is mildly convex sloping anterolingually at the
anterolingual tooth margin and curving posterolingually around the buccal margin of the
metacone. The lingual bases of the protocone and metaconule slope gently towards the
base of the crown. The lingual bases of the paracone and metacone slope more steeply into
the longitudinal tooth valley. The triangular buccal surface of the paracone 1s reduced rela-
tive to the metacone as is the paracone buccal margin. The buccal basin of the paracone is
closed and deep. The buccal basin of the metacone is comparatively shallower but remains
closed. A ridge-like stylar shelf extends along the length of the buccal tooth margin. The
relative heights of stylar cusps in descending order are as follows: stylar cusp C, B, A, D
and E. The preparacrista, postparacrista, premetacrista and postmetacrista which make up
the buccal selene of the tooth, are distinct linear crests which extend from the apices of the
paracone and metacone respectively. The lingual selene is offset posteriorly and is com-
posed of a relatively linear preprotocrista and premetaconule crista and a crescentic post-
metacrista and highly crescentic postprotocrista. The postprotocrista and premetaconule-
crista meet in the transverse median valley at a point slightly lingual to the longitudinal val-
ley. The parastyle is poorly developed. It exists as a slight swelling of the anterior cingulum
at the anterobuccal corner of the tooth. A shallow pocket is created between the anterior
cingulum and the anterior base of the paracone. Lingually, the pocket is defined by a small,
crenulate, but distinct paraconule which lies at the anterolingual base of the paracone. A
short, non-cuspate, spur-like protostyle which originates from the preprotocrista at a point
slightly lingual to the longitudinal valley, extends posteriorly along the buccal base of the
protocone, terminating opposite the protocone apex. A well developed anterolingual
paracrista connects the paraconule posteriorly to the apex of the paracone. A similarly well-
developed posterolingual paracrista extends from the apex of the paracone into the junction
of the transverse median and longitudinal valleys wherein it divides into two spurs which
become part of the crenulation pattern of the tooth.
A small, crenulate, non-cuspate neometaconule lies in the longitudinal tooth valley at
the anterolingual base of the metacone. It is connected to the metacone base by a weak
anterolingual metacrista, which originates approximately half way down the base of the
metacone. A posterolingual metacrista is absent. A crenulate, cingulum-like shelf runs along
the anterior and anterolingual base of the protocone forming a shallow pocket between the
cingulum and the protocone base. A weak anterolingually directed crest extends from the
apex of the protocone to meet the anterolingual cingulum. A similar, but deeper pocket
occupies the anterolingual base of the metaconule and is bounded by the posterior base of
the protocone anteriorly, the anterior base of the metaconule posteriorly, the junction of the
premetaconule crista and postprotocrista buccally, and a short, crescentic lingual cingulum
lingually. The lingual cingulum effectively closes off the lingual exit of the transverse medi-
an valley and is continuous with a weak crest that extends anterolingually from the apex of
the metaconule. Weak buccally directed spurs extend from the apices of the protocone and
metaconule, fading down their respective bases towards the longitudinal valley. A well
developed posterior cingulum is continuous with the postmetaconulecrista lingually and
buccally and meets stylar cusp E at the posterobuccal tooth corner.
The Mj of the holotype is poorly worn and the anterolingual tooth corner is miss-
ing. Consequently, much of the crown morphology of the trigonid has been lost. In
AMF98886 the preprotocristid is a well defined, linear crest which terminates anteriorly
in a small paraconid. The preprotostylidcristid extends anterolingually to terminate at the
base of the paraconid. Pledge (1992), in his description of the holotype Mj, suggests that
C. yorkensis differs from all other phascolarctids in having a fine anterobuccal spur of
the preprotocristid meet the preprotostylidcristid at the anterior tooth margin. Its absence
in AM F98886 suggests this is a variable feature within the species, and the prepro-
Proc. LINN. SOc. N.S.W., 117. 1997
14 GIANT RINGTAIL POSSUMS AND GIANT KOALAS
tocristid spur is most probably part of the crenulation pattern of the tooth. A well-devel-
oped posterobuccal ridge extends from the protostylid apex in AM F98886. This crest is
only vaguely discernible in the holotype. The buccal cingulum is better developed in AM
F98886. The postprotocristid meets the anterior base of the cristid obliqua at a slightly
more lingual position than in the holotype. The lingual columnar stylids of the metaconid
and entoconid are large, crenulate and well preserved in AM F98886 whereas they are
only represented by slight swellings in the holotype. The entostylid is a well-developed,
cuspate swelling on the terminus of the postentocristid at the posterolingual tooth corner
in AM F98886. AM F98887, AM F98888 and AM F98889 (M>) are morphologically
similar to but smaller than the M> of the holotype. Again, the holotype is highly worn,
the apices of all major cusps are missing and the lingual tooth margin is damaged such
that the columnar stylids are not preserved and the entostylid is missing. In contrast, all
of the Wellington Caves M> specimens show little or no wear. The columnar stylids are
large, particularly that of the metaconid. The postentocristid curves lingually to the pos-
terolingual tooth corner where it terminates in a well developed entocristid. The ento-
conulid, like in the holotype, is poorly developed. Well developed lingually directed
crests extend from the apices of the protoconid and hypoconid and terminate just prior to
reaching the longitudinal valley. These crests are not present in the holotype. The M3,
AM F98890, is largely unworn and is slightly smaller than the M3 of the holotype. The
postmetaconid, preentocristid and postentocristid are more arcuate than in the holotype.
The lingual buttresses of the metaconid and entoconid are better developed however the
buccal bases of these cusps are more greatly expanded in the holotype. The metastylid is
larger and the metastylid fold is more pronounced. The lingual faces of the protoconid
and hypoconid are more steeply sloping and the lingually directed crests associated with
these cusps are well developed (however they are absent in the holotype). Following an
analysis of variation in dentitions of the modern species, it is evident that the above men-
tioned morphological differences between the referred material and the holotype fall
within the boundaries of normal intraspecific variation.
DISCUSSION
Pseudokoala cathysantamaria 1s an enormous ringtail possum and Cundokoala
yorkensis an enormous koala. With the addition of these two to the Pleistocene of
Australia, all families of Australian herbivorous mammals are now known to have had
giants in the Pleistocene megafauna. They also significantly increase the ranks of known
arboreal megafaunal species. Apart from one modestly large koala (Phascolarctos stir-
toni Bartholomai, 1968), the only undoubted megafaunal arboreal species previously
known from the Pleistocene was Bohra paulae Flannery and Szalay, 1982, a gigantic tree
kangaroo from Wellington Caves, New South Wales.
ACKNOWLEDGMENTS
We thank the Australian Research Committee, the University of New South Wales, the Queensland
Museum, National Estate Programs (Queensland), the Riversleigh Society, Queensland National Parks and
Wildlife Service, Australian Geographic Society, Royal Zoological Society of New South Wales, Linnean
Society of New South Wales and private individuals for support that has enabled discovery and preparation of
the diverse koalas and ringtail possums of Riversleigh. Henk Godthelp, Anna Gillespie and Anne Musser (all
of the University of New South Wales) helped collect material from Wellington Caves. Armstrong Osborne
(University of Sydney) and Michael Augee (University of New South Wales) organised the opportunity to
examine the Wellington deposits. Mathew Crowther (University of New South Wales) confirmed the identifica-
tions of other Wellington mammals excavated with the tooth of Cundokoala yorkensis. Neville Pledge (South
Australian Museum) and Tom Rich (Museum of Victoria) made materials available for study. Betty Thompson
(Museum of Victoria) confirmed collection details of Portland materials. Suzanne Hand and Jenni Brammall
(University of New South Wales) read and constructively criticised a draft of this paper.
Proc. LINN. SOC. N.S.W., 117. 1997
M. ARCHER, K. BLACK AND K. NETTLE 15
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Proc. LINN. Soc. N.S.W., 117. 1997
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Functional Anatomy of the Macropodid Pes
NICOLAS BISHOP
(Communicated by M.L. Augee)
Biological Sciences, Flinders University, GPO Box 2100, Adelaide SA 5001
Bishop, N. (1997). Functional anatomy of the macropod pes. Proceedings of the Linnean
Society of New South Wales, 117: 17-50
The functional anatomy of the sthenurine pes was investigated, based on a compara-
tive study of the extant macropodines. Comparisons were made with the possum, koala, and
wombat to determine a number of plesiomorphic character states associated with the
diprotodontid pes. Dissections were made of the hind limb and pes of a number of extant
potoroids and macropodids to determine the functional nature of the actions and insertions of
the muscles, as well as the nature of the articulations. A cluster analysis was used to sort a
collection of 219 fossilised macropodid calcanea from the Pleistocene Victoria Fossil Cave
deposit, Naracoorte southeast South Australia. The morphology of the sthenurine calcanea
were compared with Macropus fuliginosus. The findings support the previously held view
that the sthenurine pes had been selectively modified for its weight bearing function during
stand-up browsing. Morphological differences between different sthenurine species suggests
that the pedal morphology could be useful in the construction of sthenurine phylogeny.
Manuscript received 1 May 1996, accepted for publication 23 Oct 1996.
KEYWORDS: Functional, anatomy, macropodid, kangaroo, foot, pes, biomechanics, mor-
phology, evolution.
INTRODUCTION
The aim of this research was to investigate the locomotion of the extinct sthenurine
kangaroos (subfamily Sthenurinae) by means of a functional analysis of their pes (hindfoot).
The locomotory adaptations of the extinct sthenurines may be inferred from a study of mod-
ern functional analogues, in this case the macropodines. The structure and function of many
aspects of the macropodine pes have been examined to varying degrees in past studies
(Parsons 1896; Windle and Parsons 1897; Barnett 1970; Lewis 1964, 1983; and Hopwood
and Butterfield 1990). Neither the form of the macropodine pes, nor that of any other ani-
mal, is dictated solely by its functional role. Form may be influenced as much by ancestry as
by functional adaptations. Unravelling these interactions is an essential aspect of functional
analysis. In this case it was necessary to determine what specific features made the pes
macropodine (the derived or apomorphic conditions) as opposed to diprotodontid (ancestral
or plesiomorphic conditions). All diprotodont marsupials, which include the Macropodidae,
share the same basic foot pattern in which the IV" digit is enlarged, sometimes equalling in
size, but usually larger than the vih digit. The vith digit is in turn larger than the syndacty-
lous II"¢ and III digits, which are again larger than the often vestigial or absent hallux or
ISt digit. This is expressed as the digital formula, [V>V>IN=U>1 .
At the basis of the concept of modern functional analogues, is the well studied con-
ception of function being reliant on form (Bock and von Wahlert 1965; Gould and
Lewontin 1979; Bock 1980, 1981, 1988, 1989; Lauder 1981; Arnold 1983; Gans 1988;
Lauder 1990; and Losos 1990). The same authors have expressed the need to use careful
consideration of all aspects of the evolution of form when ascribing particular traits to a
specific function.
To ascertain the plesiomorphic diprotodontid pes structure from which the
Proc. LINN. Soc. N.S.w., 117. 1997
18 FUNCTIONAL ANATOMY OF THE MACROPODID PES
macropodine pes has evolved, it was necessary to examine the characteristics of several
different marsupial families. Evidence from studies of craniodental characters place the
Macropodidae close to, but derived from, the Potoroidae. Together these two are closely
related to, but derived from, the Phalangerids (possums), (Dollo 1899, Bensley 1903,
Raven and Gregory 1946, Archer 1984, and Flannery 1983). The superfamily
Macropodoidea is broken up into two main families, the Macropodidae and Potoroidae.
The muscular anatomy of the hindlimb and pes of the macropodines has previously
been examined by Parsons (1896), Windle and Parsons (1897), Craven (1971), Adnams-
Hodges (1988), and most recently and comprehensively by Hopwood and Butterfield
(1990). Only Parsons (1896) and Lewis (1983) have investigated the anatomy of the
binding ligaments of the pes. The osteological remains of the macropodine foot and the
nature of the articulations have also been dealt with to various degrees by Owen (1875);
Parsons (1896); Windle and Parsons (1897); Barnett (1970); Lewis (1964, 1983); and
Hopwood and Butterfield (1990).
The locomotion of the macropodids, initially documented by Muybridge (1887)
has been well studied, particularly by Badaux (1965), Windsor and Dagg (1971),
Marshall (1974), Bennett (1987), Biewener and Blickhan (1988), and Baudinette (1994).
Macropodines progress by a ricochetal saltatory mode of locomotion, where both of the
hind-feet are placed on the ground simultaneously. Weight is transmitted through the
tibia to the astragalus, the calcaneum, through the cuboid, then through to the elongated
fourth metatarsal. Windsor and Dagg (1971) in their study of nineteen macropodine
species were able to identify four gaits: 1) the slow progression - pentapedal (four limbs
and tail), used mainly during grazing; 2) the walk - the pairs of the limbs are asynchro-
nously placed in contact with the substrate, and expressed only in Dendrolagus the tree
kangaroo; 3) the quadrupedal bound - the use of the hind feet followed by the forefeet,
exhibited only in Setonix, Dendrolagus, Petrogale and the Potoroidae; 4) the bipedal hop
- he fastest gait, with synchronous placement of the hind feet on the ground.
Independent studies by Marshall (1974), and previously Badaux (1965), charac-
terised macropodine locomotion as hopping or bipedal ricochetal saltation, where the
hind feet are synchronously placed in contact with the ground. Despite macropodines
possessing at least four gaits, it seems that the general elongated form of the macropo-
dine locomotor apparatus has been modified primarily in relation to the fastest gait, the
bipedal hop. This is supported by evidence associated with the energetic efficiency of
this gait at high speeds (Dawson and Taylor 1973, Alexander and Vernon 1975, Cavagna
et al. 1977, Baudinette et. al. 1992, Baudinette 1994), which can be primarily explained
by the physical return of stored energy in the hindlimb (Alexander and Vernon 1975,
Alexander 1984, and Bennett and Taylor 1995).
Based on a study of cranial and dental characters, Raven and Gregory (1946)
examined the adaptive branching of the kangaroos and its relation to habitat. The pes can
also be used in a similar function as eluded to by Windsor and Dagg (1971). The
macropodines have diversified to occupy a wide variety of habitats ranging from open
plains (Macropus rufus) to tree top canopies (Dendrolagus). An examination of the var-
ied pedal morphology of the macropodines from segregated habitats should provide an
excellent source of definition for the functional nature of the derived features.
Wells and Tedford (1995) noted some major differences between the sthenurine kan-
garoos, and the extant (modern) macropodines. The first major difference is in the general-
ly bulkier nature of the animal; the skull and teeth being adapted for tough feed browsing,
rather than grazing which predominates in the modern macropodines. The structure of the
shoulder facilitated raising the arms above the head, with the associated long digits of the
manus (hand) enabling it to reach higher foliage. Structure of the spine and tail vertebrae
suggest adaptations to standing upright, also for procuring food. These authors also note
that the hindfeet are functionally one-toed or monodactylous, with further reduction of the
second, third, and fifth digits differing from the modern generalised macropodine form.
Proc. LINN. SOc. N.S.W., 117. 1997
N. BISHOP 19
MATERIALS AND METHODS
All of the fossil bones used in this study were excavated from the “Victoria Fossil
cave” at Naracoorte from 1972 to the 1988. Measurements were also taken on various
reference skeletons from extant species in Flinders University and South Australian
Museum collections. The specimens are listed in Table 1.
TABLE |
The reference skeletons examined from the South Australia Museum (M) and Flinders University Reseacrh
(FUR) collections.
Specimen Reference
POTOROIDAE Hypsiprymnodon moschatus M11940
Aepyprymnus rufescens FUR
Bettongia lesseur FUR O11
Bettongia penicillata M8286
MACROPODIDAE Macropus fuliginosus FUR
Macropus rufus FUR 001
Macropus eugenii FU 010
Macropus rufogriseus M16370
Macropus greyi M2121
Macropus parma M7191
Macropus irma M16489
Macropus robustus M3695
Macropus parryi M14103
Macropus dorsalis M7967
Dorcopsis sp. M13754
Petrogale lateralis M12555
Setonix brachyurus M14102
Wallabia bicolor M16469
Dendrolagus bennettianus M5530
PHALANGERIDAE Pseudochirus peregrinus FUR 004
VOMBATIDAE Lasiorhinus latifrons FUR 006
PHASCOLARCTIDAE Phascolarctos cinereus
FUR 009
Muscles, tendons and ligaments
The hind limbs were removed from a number of kangaroo carcasses for the pur-
pose of dissection. Whole frozen specimens of one female western grey (M. fuliginosus)
and one female red kangaroo (Macropus rufus) were provided by the South Australian
Proc. LINN. SOc. N.S.W., 117. 1997
20 FUNCTIONAL ANATOMY OF THE MACROPODID PES
National Parks and Wildlife Service. One Tammar wallaby (Macropus eugenii), one bur-
rowing bettong (Bettongia lesueur) and one rufous bettong (Aepyprymnus rufescens)
from the frozen specimen collection at Flinders University were dissected.
All specimens had been frozen soon after death and stored in a freezer room held
at -15°C. Two to three days prior to their dissection they were transferred to a refrigerat-
ed room held at 4°C. The specimens remained in the refrigerated room for the duration,
to minimise deterioration of the flesh. All dissections were carried out using a standard
dissection kit. After all of the flesh was dissected away the limbs of the specimens were
macerated, cleaned and bleached so that they could be illustrated and examined for mus-
cle and ligament scars.
The measurements taken on the calcaneum are displayed in Fig. 1. The terms used
to describe the features of the pes follows Murray (1995). Figure 2 shows the anatomical
terms used in the descriptive anatomy.
Statistical analysis
Twelve measurements were made on the calcanea of both the extinct and extant
taxa. A cluster analysis was performed on this data matrix to determine the natural
groupings of the specimens measured.
The statistical program SPSS (1989) was used to cluster the fossil bones into nat-
ural groupings based on the measurements taken. Recognition of the distribution pat-
terns of operational taxonomic units (OTU) and groups of OTU’s (taxa) were carried
out in a hyper dimensional (12 dimensions) space (phenetic A space), where a pattern
is any discernible property of the distribution and groups of OTU’s in A space. I have
chosen a cosine measure of similarity. The use of cosine as a similarity measurement is
particularly useful in the measurement of shape (Sneath and Sokal 1973). This stan-
dardises size so that measurement of similarity is on shape only. The agglomerative
clustering technique was used to cluster the OTU’s and is the most widely accepted
and tested method. There are several methods available to group the specimens, how-
ever the most widely accepted, and also default setting for the SPSS program, is the
sequential, agglomerative, hierarchical, non-overlapping clustering method (SAHN).
To summarise the differences, a principal component analysis was carried out on
the same data set. Principle components analysis measures the variance for the entire
group and then displays the variance in the form of major components.
RESULTS
Comparative Anatomy and Cladistic Analysis of the Marsupial Pes (Tables 2-4)
Anatomy of the crus and pes of the western grey kangaroo, Macropus fuliginosus
Osteology
Separate bones of the crus and pes include: tibia, fibula, seven tarsal bones, four
metatarsals, twelve phalanges, a plantar sesamoid, four metatarsophalangeal sesamoids
(Figs 3 and 4).
i) The crus: (between knee and ankle) comprising of tibia and fibula: On the tibia
the condyles are located on the posterior half of the head of the tibia. The tibia articulates
with the fibula along the posterolateral aspect of the lateral condyle of the tibia. The tibia
is greatly expanded anteroposteriorly compared with its transverse width. Proximally the
tibia is expanded, with the shaft roughly triangular in cross section as described by Owen
(1874-77). Distally the shaft is circular to ovoid at its extremity, expanding into the
medial malleolus. The proximal lateral surface of the shaft is deeply concave to receive
the extensor muscle mass of the crus.
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 2]
4 ss
aS SO
Figure 1. a-The length of the calcaneum from the epiphysial junction to the line of transverse axis of the astra-
galus: b- The length of the calcaneum from the epiphysial articulation to the distal portion of the dorsolateral
cuboid facet on the calcaneum: c- The width of the dorsal portion of the tuber calcanei of the calcaneum mea-
sured transversely at the posterior region of the lateral process and the posterior region of the sustentaculum
tali: d- Width across the pivot point, or line of transverse axis of the astragalus: e- The width of the plantar sur-
face of the calcaneum measured at the most posterior aspect of the tuber calcanei, not including the epiphysis if
present: f- Plantar surface length, measured from the epiphysial arthrosis on the plantar surface to the trans-
verse plantar sulcus on the calcaneum or when not present, to the distal portion of the plantar tuberosity on the
calcaneum: g- The prominence of the dorsolateral cuboid facet on the calcaneum, measured from the face of
the dorsomedial cuboid facet on the calcaneum to the most distal portion of the dorsolateral cuboid facet on the
calcaneum: h- The height of the calcaneum measured dorsoventrally at the most posterior aspect of the tuber
calcanei, not including the epiphysis if present: i- The surface area of the dorsomedial cuboid facet on the cal-
caneum: j- The surface area of the dorsolateral cuboid facet on the calcaneum: k- The surface area of the ven-
tromedian cuboid facet on the calcaneum: |- The angle of transverse axis of the astragalus measured from the
central axis of the calcaneum.
Proc. LINN. SOC. N.S.W., 117. 1997
2D FUNCTIONAL ANATOMY OF THE MACROPODID PES
TABLE 2
The characters which were determined to be plesiomorphic.
Character State
] Weak and asymmetrical ridges
2 Malleolar fossa is situated posteromedially, and not distinct
3 The medial trochlear crest is greater in length than the lateral trochlear crest
4 Large scar for the posterior talocalcaneal ligament
5 Long astragalar head and neck with no distinct notch for the trochlear bursa
6 The lateral trochlear crest is aligned with the lateral border of the navicular facet on the astragalus
7 The medial trochlear crest aligns with the lateral border of the navicular facet on the astragalus
8 The navicular facet is broad, with its longitudinal axis transecting the lateral trochlear crest of the
astragalus
9 Transversely wide sustentaculum tali which tapers medially
10 Uninformative
11 The sustentaculum tali is deflected anteromedially, and the base of the sustentaculum tali is rounded
posteriorly
12 The articular surface for the astragalus on the calcaneum is continuous transversely, and shallowly
convex anteroposteriorly
13 The lateral facet for the astragalus on the calcaneum is higher than the medial facet for the astragalus
Derived characters of other diprotodontids:
5 The wombat possesses both a long astragalar head and neck and a small notch for the trochlear bursa
11 In the koala, the base of the sustentaculum tali is straight, nearly horizontal
14 Single smooth surface distally for articulation with the cuboid in other diprotodontids
15 No transverse plantar sulcus in other diprotodontids
16&17 Single articulating surface for the cuboid in other diprotodontids
TABLE 3
The shared derived characters of Hypsiprymnoden and Macropus.
Character State
5) cree The trochlear ee sqmmenieal and wall rounded
6 The lateral trochlear crest is aligned with the lateral portion of the dorsolateral facet (of the caleaneum)
when the astragalus is articulated
7 The medial trochlear crest aligns with the median portion of the navicular facet on the astragalus
9 Comparatively transversely narrow astragalus
14 The dorsomedial and dorsolateral facets (for the cuboid) are equal in height in distal aspect
16 Presence of a groove separating the dorsomedial and ventromedian facets (for the cuboid) on the
calcaneum
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 23
TABLE 4
The derived or apomorphic characters of the generalised macropodine pes.
Character State
There is a distinct notch on the transversely wider medial malleolar fossa
2
3 The parallel trochlear ridges are angled more obliquely
4 The scar for the posterior talocalcaneal ligament is much reduced
5 The development of a distinct notch for the trochlear bursa
8 The navicular facet is broader, and its longitudinal axis transects the medial trochlear crest
10 The tuber calcanei is relatively narrow and elongated anteroposteriorly
11 The sustentaculum tali is deflected anteromedially, and curved posteroventrally
WW There are two separate lobate facets (on the calcaneum) for articulation with the astragalus
13 The medial and lateral facets (for the astragalus) on the calcaneum are equal in height
15 The transverse plantar sulcus is anteroposteriorly narrow
17 The cuboidocalcaneal step is short but distinct
The fibula
is a long and slender bone, with the head being the most expanded part, anteropos-
teriorly. It articulates with the transverse fibular groove on the posterolateral aspect of
the tibia. The fibula shaft is thickest proximally, with the lateral surface, convex and the
medial surface concave to articulate with the lateral aspect of the tibia, as well as provid-
ing area for muscle insertion in proximal region. The distal epiphysis is expanded into
the lateral malleolus (Hopwood and Butterfield 1990).
11) The Pes consists of a number of separate bones:The tarsus consists of seven
separate tarsal bones plus one sesamoid (Hopwood and Butterfield 1990): calcaneum,
astragalus (talus), and Metatarsal IV make up the bulk of the tarsus.
The astragalus
(talus) bears a large articulation dorsally with the distal end of the tibia. The bone
is subdepressed and triangular , with the base turned forward (Owen 1874-77). In medial
or lateral view it is arched, convex dorsally. The dorsal surface is formed mainly by the
trochlear articulation with the tibia, which is convex anteroposteriorly and concave trans-
versely. Further, the dorsal surface is divided into medial and lateral portions by the
respective medial and lateral trochlear grooves, providing facets for the lateral and medi-
al malleoli. Ventrally the astragalus is concave anteroposteriorly, forming the articular
surface for the calcaneum. It is divided into a lateral and medial articulation site, corre-
sponding to articulation sites on the calcaneum. The lateral surface of the head of the
astragalus also articulates with a small area on the dorso medial portion of the calca-
neum. Distally, the astragalus articulates with the navicular bone as well as a small area
on the posteromedial portion of the cuboid.
The calcaneum
articulates dorsomedially with the astragalus, dorsolaterally with the fibula, and
distally with the cuboid. There is a roughened flat surface on the plantar aspect for the
insertion of the large plantar calcaneo cuboid ligament. The calcaneum is roughly trian-
gular in cross section, flattened and broad plantarly. The dorsal surface of the tuber cal-
canei has a smooth surface with a relatively narrow longitudinal ridge extending posteri-
orly. The sustentaculum tali is narrow transversely but deep dorsoplantarly, and is curved
Proc. LINN. Soc. N.S.W., 117. 1997
24 FUNCTIONAL ANATOMY OF THE MACROPODID PES
Median Plane
Sagittal Planes
Figure 2. Shows the anatomical and directional terms and planes of the body (Adapted from Leach, 1993)
posteriorly through plantarly. It bears a groove for the flexor digitorum longus. Laterally,
the fibular condyle bears an articular facet for the lateral malleolus of the fibula on its
posterolateral aspect. Dorsally, the lateral facet for the astragalus is high and rounded,
proportioned equally transversely and anteroposteriorly. It is separated from the concave
medial facet by a small longitudinal ridge. The ridge of the medial facet sits slightly
higher and more anterior to the posterior border of the lateral facet. The fossa for the
anterior proximal process of the astragalus is shallow and semicircular. There is a small
triangular facet on the dorsomedial aspect of the calcaneum for articulation with the lat-
eral aspect of the neck of the astragalus. Distally, the surface of the dorsomedial facet is
transversely broad, rectangular and strongly convex dorsoplantarly. The dorsolateral
facet is narrow transversely, but longer dorsoplantarly and narrowing plantarly. This
facet is stepped and protrudes more anteriorly than the dorsomedial facet. Ventrally, the
dorsolateral facet merges into the ventro median facet, which is of similar size to latter,
but broader transversely than dorsoplantarly, and is of sub triangular shape, but with the
most plantar aspect not contributing to the plantar surface.
The navicular
proximally, is concave, articulating with the corresponding convex head of the
astragalus. Laterally, it articulates with the medial surface of the cuboid. It is convex and
articulates distally with the entocuneiform and ectocuneiform.
The entocuneiform
articulates proximally with the navicular, distally with the 1194 metatarsal and
mesocuneiform, laterally with the cuboid, and medially with the ectocuneiform
(Hopwood and Butterfield 1990).
The mesocuneiform
articulates proximally with the ectocuneiform, distally with the 1194 metatarsal,
dorsally with the [14 metatarsal, and is the smallest tarsal bone.
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 25
Figure 3. View of the ligaments of the left pes of Macropus fuliginosus . TCL- tibiocalcaneal ligament ;
PTTL- posterior tibiotalar ligament ; MEN- meniscus ; PTFL- posterior talofibular ligament ; PCFL- posteri-
or calcaneofibular ligament ; ACFL- anterior calcaneofibular ligament ; AHM- anterior horn of the meniscus ;
PCCL- plantar calcaneocuboid ligament ; LCCL- lateral calcaneocuboid ligament ; MCCL- medial calca-
neocuboid ligament ; TNL- talonavicular ligament ; TCL- talocuboid ligament ; PTCL- posterior talocalcaneal
ligament ; PMTCL- posteromedial talocalcaneal ligament ; LCT- ligamentum cervicis tall.
Proc. LINN. Soc. N.S.W., 117. 1997
26 FUNCTIONAL ANATOMY OF THE MACROPODID PES
The ectocuneiform
articulates proximally with the navicular, distally with the mesocuneiform and md
metatarsal, medially with the entocuneiform, and laterally with the cuboid and Ivt
metatarsal, extending backward, beyond and overlapping the entocuneiform (Owen
1874-77).
The cuboid
articulates proximally with the calcaneum accommodating its three facets, distally
with IV" and vt metatarsals, and medially with the navicular and ectocuneiform.
Metatarsal I
Absent
Metatarsal II and III
Metatarsal II articulates proximally with entocuneiform and mesocuneiform, and
distally with the first phalanx of digit Il. Metatarsal III articulates proximally with meso-
cuneiform and entocuneiform, and distally with the first phalanx of digit III. Both
metatarsals II and III are reduced greatly in width but not in length. In the middle of the
tarsus both lie on the plantar aspect of metatarsal IV, then curve dorsomedially in the dis-
tal portion of the metatarsus (Hopwood and Butterfield 1990).
Metatarsal IV
articulates proximally with the cuboid, and proximomedially with the ecto-
cuneiform. It articulates medially with metatarsal III, laterally with metatarsal V, and
plantarly with the tarsal sesamoid bone (Hopwood and Butterfield 1990). Distally
metatarsal IV articulates with the first phalanx of digit IV and the medial and lateral
metatarsophalangeal sesamoid bones. The dorsal surface is convex transversely, most so
at the most proximal part where it articulates with the cuboid. The plantar surface is
slightly concave transversely, mainly in the distal two thirds, but with a deep tuberous
keel proximally present as a thick ridge. The proximal plantar surface also provides a
site for the attachment of the plantar sesamoid, which is grooved for accommodation of
the flexor digitorum longus tendon. The distal articulation is convex dorsoplantarly but
nearly flat transversely. Ventrally on the distal articular surface is a median ridge form-
ing two concave surfaces for the trochlear articulation of the first phalanx of the
metatarsal IV.
Metatarsal V
articulates proximally with the cuboid, and medially with the lateral surface of
metatarsal IV. It articulates distally with the first phalanx of digit V and also plantarly
with the metatarsophalangeal sesamoid bones of digit V. A significant portion of
metatarsal V is in contact with the ground. Proximally this bone possesses a sigmoidal
shape, curving up most proximally at its articulation with the cuboid. The rest of the
bone is curved in the opposite direction, very greatly curved concavely. Proximally the
bone is roughly triangular in cross section. The lower border is thicker than the upper,
and primarily in the proximal portion. In plantar aspect metatarsal V is also curved out
laterally so that the distal end of this bone lies ventrolaterally to metatarsal IV. The distal
articulation with the first phalanx of digit V is convex dorsoplantarly.
Digits
There are three digits corresponding to each of the metatarsals, and the size of
these digits conforms well with the size of the metatarsal carrying that digit. The
metatarsal IV is the longest digit with the distal digits of metatarsal V extends as far as
the articulation between the first and second digits of metatarsal IV.
Proc. LINN. Soc. N.S.W., 117. 1997
N. BISHOP 27
The ligaments of the pes
In the majority of mammals including humans, the ankle consists of only one joint.
It is noted that in macropods the ankle has been modelled into two ankles (Lewis 1980).
The first joint is considered the articulation of the tibia with the astragalus and the fibula
with the calcaneum, with the second joint (sub talar) being formed between the astra-
galus and the calcaneum. The Nomina Anatomica text suggests that the ankle joint is the
talocruralis, i.e. the joint between the tibia and the astragalus. While this joint exists in
the macropods, the fibula also articulates with the calcaneum. The articulation of the
tibia on the astragalus is a “hinge-like” trochlear joint convergent in form and function
with the eutherians (Lewis 1980).
i) Articulatio tarsocruralis (tibiotalar/calcaneofibular) (see Fig. 3): The broader lat-
eral part of the tibial surface articulates directly with the large trochlear groove on the
astragalus. This area is separated from the medial articulation by the medial trochlear
ridge. The well defined astragalar depression accommodates the terminal articular knob
on the tibia. The lateral trochlear ridge separates the trochlear groove from the lateral
surface of the astragalus where the medial surface of the fibula articulates.
Medially, there is a ligamentous wall, common to congruent joints. The posterior
tibiotalar ligament arises from a long ridge, extending longitudinally on posteromedial
aspect of the astragalus. More superficial to this and crossing over the tibiotalar ligament,
is the tibiocalcaneal ligament which passes down to attach to the tuberosity on the sus-
tentaculum tali. This is in close proximity to the talo-navicular ligament passing from the
medial aspect of the neck of the astragalus to a dorso-lateral position on the navicular. A
small cartilaginous meniscus intervenes on this surface in between the fibula and the
astragalus, but does not extend up in between the tibia and fibula (Lewis 1980).
Anteriorly, the meniscus continues forward between the fibula and the astragalus,
terminating as a fibrous horn, and recurves to attach to the calcaneum. Passing from the
fibula to the astragalus, protruding from underneath the meniscus is the posterior
talofibular ligament which intervenes between the fibula and the exposed projection of
the articular facet on the calcaneum. Also protruding from the astragalus near the astra-
galar neck, the talocuboid ligament divides into two components; one band transversely
forming a strap through which the tendons of the flexor muscles pass, and another band
over the cuboid to insert into its lateral surface, where it shares its insertion with a small
lateral calcaneocuboid ligament. As with the medial side of the tibia, two ligaments, the
posterior and anterior ligaments, cross and bind the fibula and calcaneum, both with ori-
gins on the lateral projection of the calcaneum and insertions on their respective diago-
nally opposite anterior and posterior regions on the distal end of the fibula. On the plan-
tar surface there are two main ligaments binding the calcaneum to the more distal tarsals
and metatarsals (Lewis 1980). Crossing from the medial side posteriorly on the calca-
neum to an insertion into the cuboid and base of metatarsal IV and V on the ventro-later-
al aspect is the large plantar calcaneocuboid ligament. Medially is a comparatively small-
er calcaneocuboid ligament originating in the base of the sustentaculum tali and passing
to an insertion into the navicular, cuneiform bones and metatarsal II and III.
i1) Talocalcaneocentralis (subtalar joint): This is comprised of the articulation of
the astragalus on the medial portion of the calcaneum. There are two main articulating
surfaces on the plantar aspect of the astragalus corresponding to the lateral and medial
articulating surfaces on the calcaneum. Both surfaces have a predominantly transverse
axis, are concave on the astragalus and are convex on the calcaneum (Lewis 1980).
Two main ligaments tightly bind the astragalus to the calcaneum, the anterior and
posterior talocalcaneal ligaments. The anterior talocalcaneal ligament is the homologue
of the ancestral ligamentum cervicis tali. It extends from the neck of the astragalus to a
large insertion into the median portion of the calcaneum (Lewis 1983). The large posteri-
or talocalcaneal ligament extends from a large area of insertion on the astragalus to a
similar sized tuberosity in the posterolateral region of the articulating part of the calca-
Proc. LINN. Soc. N.S.W., 117. 1997
28 FUNCTIONAL ANATOMY OF THE MACROPODID PES
neum. There is also a small ligament extending between the postero-medial process of
the astragalus to the corresponding postero-medial area on the calcaneum. This ligament
is not described in any text, and is probably a second fascia of the main posterior talocal-
caneal ligament. Similar remodelling of the subtalar joint such as seen in macropods can
also be found in the precursors of the artiodactyls (Lewis 1983).
The muscles and tendons of the crus and pes of the western grey kangaroo, Macropus
fuliginosus
The complete descriptions of the muscles of the crus of the eastern grey (Macropus
giganteus) have been succinctly described in a previous study, (Hopwood and Butterfield
1990) and will not be dealt with here.
Comparative morphology of modern Macropodines
The muscles and tendons of the hindlimb and pes
The muscles and ligaments of a number of modern macropodid and potoroid forms
were examined to determine whether there were any differences in the origins and particu-
larly insertions of the muscles, and to see whether these differed significantly from the
form of M. fuliginosus. The insertion sites of the tendons did not vary between the different
macropod species examined in this study. It was noted however that there was a correlation
between the size of the tendon entering an insertion site and the size of the resulting scar.
The comparative osteology of Macropus fuliginosus, Macropus rufus, and Dendrolagus
bennettianus
To ascertain some of the features which may be related to habitat in the modern
kangaroos, I have chosen to describe two species from two different habitats using
Macropus fuliginosus as a comparative model.
i) Comparative anatomy of the astragalus: Dorsally in Dendrolagus bennettianus the
astragalus is subdepressed and rectangular in form, greater transversely than anteroposte-
riorly, compared with M. fuliginosus which is slightly more triangular in form, due largely
to the angle of the lateral trochlear crest. Macropus rufus is the most triangular of the three
specimens with the medial trochlear crest greatly elongated posteriorly. The dorsal surface
of the astragalus in the three specimens is formed mainly by the trochlear-like articulation
with the tibia. The dorsal surface is divided into three portions by the two-medial and lat-
eral trochlear crests providing facets for the lateral and medial malleoli. The astragali of
the specimens are curved convexly anteroposteriorly, greatest in Dendrolagus and least
curved in M. rufus in keeping with the proportions of the relative calcanea. The trochlear
of the astragalus is also curved concavely transversely, deepest in Dendrolagus and most
shallow in M. rufus compared with M. fuliginosus. The comparative height of the medial
and lateral trochlear crests varies between the three specimens. In M. fuliginosus and M.
rufus the medial trochlear ridge is higher than the lateral trochlear ridge viewed in distal
aspect, compared with D.bennettianus where these ridges are equal in height.
Ventrally the astragalus is concave anteroposteriorly, forming the articular surface
for the calcaneum, being approximately equal depth in all three specimens. The trans-
verse width of the articular facets is greater in D. bennettianus than M. fuliginosus which
is in turn greater than M. rufus where the facets are longer anteroposteriorly and narrow-
er transversely. The two facets for the calcaneum on the astragalus are more similar in
shape in M. rufus and M. fuliginosus compared with Dendrolagus, where in the former
two, the lateral facet is roughly circular, compared with the triangular form of the latter.
Also in Dendrolagus, the medial facet is sub triangular compared with the transversely
wide rectangular form of the medial facet for the calcaneum in M. rufus and M. fuligi-
nosus. A small area of the lateroplantar portion of the head of the astragalus also articu-
lates with a dorsomedial portion of the calcaneum.
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 29
Medially, the articulation for the medial malleolus varies slightly between the three
specimens. In Dendrolagus there is an ovoid pit on the medial malleolus compared with
a more circular pit in M. fuliginosus and M. rufus. The ridge on the medial aspect for the
posterior tibiotalar ligament is longer in M. rufus than M. fuliginosus because of the elon-
gated portion of the medial trochlear ridge. The site for the attachment of the posterior
tibiotalar ligament in Dendrolagus 1s more angled, giving a greater area of attachment for
the ligament dorsally compared with the nearly flat ridges of M. fuliginosus and M. rufus.
The area for attachment of the anterior tibiotalar ligament is smallest in Dendrolagus, but
is borne in a deep groove which lies medially and plantarly compared with M. rufus and
M. fuliginosus where the site for attachment is predominantly on the medial surface, and
also dorsally as illustrated by a deep pit in M. rufus.
Ventrally in medial aspect, the astragalus is curved concavely, most greatly in
Dendrolagus and least curved in M. rufus. The other main distinction is with the articula-
tion on the astragalus for the cuboid and navicular. In Dendrolagus the articular surface
is curved gently convexly and runs nearly transversely, terminating medially compared
with M. rufus and M. fuliginosus where this articular facet is directed dorsoplantarly. The
largest articulation on the astragalus for the cuboid is in M. rufus where this articulation
encroaches on the anterior or distal face of the head of the astragalus, so that the area on
the astragalus for the navicular is reduced and narrow dorsally. This facet is also deflect-
ed more plantarly than in Dendrolagus and M. fuliginosus.
i1) The comparative anatomy of the calcaneum: The tuber calcanei of the calca-
neum of M. fuliginosus and M. rufus is triangular in cross-section, being flattened and
broad plantarly compared with D. bennettianus which is much shorter and broader and
ovoid in cross-section, reminiscent of possums and koalas. The dorsal surface of the
tubercalcanei is formed by two surfaces which converge dorsally into a longitudinal
ridge which extends posteriorly. In D. bennettianus this ridge is very flat and broad, nar-
rowing posteriorly. In M. fuliginosus this ridge is relatively narrow, but not as narrow as
in M. rufus. There is considerable difference in the width of the sustentaculum tali of
these three macropods. In Dendrolagus the sustentaculum tali is very broad transversely
and pointed medially and shallow dorso-plantarly, compared with M. fuliginosus and M.
rufus where the sustentaculum tali is very narrow, particularly so in M. rufus.
Corresponding with the astragalus, the anteroposterior length of the sustentaculum tali is
shorter in D. bennettianus than M. fuliginosus which is shorter than the very elongated
form in M. rufus. The sustentaculum tali is deepest and most curved in M. fuliginosus. In
D. bennettianus the plantar process of the sustentaculum tali extends further medially
and nearly contacts the ground, compared with the slightly concave nature of the susten-
taculum tali dorsoplantarly in the other two specimens. The sustentaculum tali also bears
the tuberosity for the ligamentum cervicis tali on the dorsal surface and is larger in M.
rufus than M. fuliginosus compared with D. bennettianus, where the deep groove for this
ligament extends posteriorly on the medial surface. In D. bennettianus the sustentaculum
tali bears a very shallow and broad groove for the tendon of flexor digitorum longus
compared with the relatively deep groove of M. fuliginosus and M. rufus, and is more
steeply inclined in the former. Laterally, the fibular condyle exhibits similar variation.
The calcaneum has least area of contact for the fibula in Dendrolagus, and greatest in M.
rufus. In D. bennettianus the lateral malleolar condyle is angled more obliquely than in
the other macropods, as in the case with its medial malleolar condyle. Only M. fuligi-
nosus and M. rufus bear a true condyle for the fibula, posterolaterally in the former and
laterally in the latter. In M. fuliginosus and M. rufus there is an area excavated in the
body of the calcaneum underneath the lateral projection which is not present in D. ben-
nettianus. Dorsally the lateral border of the most distal portion of the calcaneum is
directed medially in M. fuliginosus and M. rufus compared with D. bennettianus, where
this section continues in the axial longitudinal plane of the foot.
There are two facets on the calcaneum for the astragalus. The lateral facet is nar-
Proc. LINN. Soc. N.S.W., 117. 1997
30 FUNCTIONAL ANATOMY OF THE MACROPODID PES
rowest in M. rufus, almost circular, slightly wider in M. fuliginosus, and much broader
transversely in D. bennettianus which narrows medially. This convex facet is separated
from the concave medial facet in M. fuliginosus by a small longitudinal ridge, compared
with D. bennettianus and M. rufus where it is nearly continuous between the two facets.
In M. rufus and D. bennettianus the ridge of the medial facet is higher than the lateral
facet. This ridge, which is curved in M. fuliginosus and M. rufus, is straight and runs
transversely in D. bennettianus.
The fossa for the anterior proximal process of the astragalus which 1s a small circu-
lar pit in M. fuliginosus, is more ovoid anteroposteriorly in M. rufus and divided by a
small ridge, and is not discernible in D. bennettianus as this portion of the calcaneum is
flat and converges with the lateral border. The corresponding fossa in D. bennettianus is
on the anterior face of the lateral facet for the astragalus. In M. rufus and M. fuliginosus
there is a small triangular facet on the dorsoanterior medial portion of the calcaneum for
articulation with the lateral border of the neck of the astragalus.
Distally, the dorsomedial facet on the calcaneum for the cuboid is transversely
broad and rectangular in M. fuliginosus, equally broad and deep in M. rufus and triangu-
lar in D. bennettianus. This facet is convex dorsoplantarly, only slightly in D. bennet-
tianus, moderately convex in M. fuliginosus, and strongly convex in M. rufus. The dorso-
lateral facet on the calcaneum for the cuboid in M. fuliginosus and M. rufus is narrower
transversely and elongated dorsoplantarly. However in D. bennettianus this facet is trans-
versely broad, and triangular in shape, narrowing laterally. This facet is convex dorso-
plantarly and concave transversely only slightly in D. bennettianus, moderately in M.
fuliginosus and strongly in M. rufus. There is a step between the dorsolateral and dorso-
medial facets on the calcaneum for the cuboid which is subtle in D. bennettianus and lies
oblique to the axis of the foot, slightly more acute in M. fuliginosus and in line with the
axis of the foot in M. rufus. The dorsolateral facet merges into the ventromedian facet for
the cuboid on the calcaneum, which is circular and slightly concave in M. rufus, circular
and slightly convex in M. fuliginosus and barely distinguishable in the flattened surface
of D. bennettianus. These three aforementioned facets in the three specimens are separat-
ed differently from contact with the ground by a sulcus. The plantar portions of the dor-
somedial and ventromedian facets are in contact with the ground in D. bennettianus and
M. fuliginosus, compared with M. rufus where the facets are separated by a large, deep
sulcus.
iii) Comparative anatomy of the cuboid: Dorsally, the cuboid is greater in length
medially than laterally in all of the specimens. The medial border is concave slightly in
M. rufus and moderately in both M. fuliginosus and D. bennettianus. The fossa for the
dorsolateral facet of the calcaneum for the cuboid is visible in dorsal view. This fossa is
obscured in M. rufus, partly visible in M. fuliginosus and entirely visible in D. bennet-
tianus. The step between the dorsolateral and dorsomedial facets of the calcaneum for
the cuboid is also apparent on the proximal surface of the cuboid, and when viewed dor-
sally this angle is in line with the long axis of the foot in M. rufus, slightly oblique in M.
fuliginosus and very oblique in D. bennettianus. The proximal portion on the medial side
in M. rufus is deflected posteromedially to articulate with the astragalus, compared with
D. bennettianus and M. fuliginosus where there is only small surface for articulation for
the astragalus on the medial side. Also in dorsal view, none of the distal articulating
facets can be seen in D. bennettianus.
Medially, the cuboids of M. rufus and M. fuliginosus are vastly different from that
of D. bennettianus, the latter which 1s rhomboidal in appearance, while in the former pair
they are rectangular, deep and anteroposteriorly compressed. The cuboid as in the calca-
neum is greatly compressed dorsoplantarly. The fossa for the dorsomedial facet of the
calcaneum can also be seen in cross-section, and it is greatly concave in M. rufus, slight-
ly less concave in M. fuliginosus and only gently concave in D. bennettianus. Also
prominent medially in D. bennettianus is a deep sulcus bearing the large cubonavicular
Proc. LINN. SOC. N.S.w., 117. 1997
N. BISHOP 31
and mesocuneiform ligament. Medially in D. bennettianus the medial plantar process
forms part of the plantar surface of the foot, being different from M. fuliginosus and M.
rufus where the medial plantar crest is raised off the plantar surface to make room for the
large tendon of flexor digitorum longus.
Anteriorly in medial aspect in D.bennettianus the cuboid is expanded dorsally
where it is concave, and convex plantarly. In contrast, in M. fuliginosus and M. rufus the
anterior profile of the cuboid is concave dorsally, convex in the median section, then
concave again plantarly. Laterally, the cuboid is longest anteroposteriorly and shallowest
dorsoplantarly in D. bennettianus. The lateral plantar crest is expanded anteroposteriorly
in M. rufus and M.fuliginosus, compared to the very short lateral plantar crest of D. ben-
nettianus. All three specimens have a deep tuberous sulcus for the insertion of the tendon
of the peroneus longus.
Ventrally the most prominent feature of M. fuliginosus and M. rufus is their large
ovoid lateral plantar process, separated from a smaller and shorter medial plantar process
by a sulcus running longitudinally, the path of the ligament attaching the medial side of
the cuboid to the base of the fourth and fifth metatarsals. In D. bennettianus the two
plantar processes, which are of equal sizes, both make contact with the ground and are
separated by an oblique sulcus running posterolateral to anteromedial region of the
cuboid.
Distally, the cuboid bears three facets; the largest articulating with the fourth
metatarsal, a smaller circular facet in M. fuliginosus and M. rufus or small triangular
facet in D. bennettianus articulating with the plantar crest of the fourth metatarsal, which
is a small rectangular facet in M. rufus and M. fuliginosus and a semicircular facet in D.
bennettianus. The facet for the plantar crest of the fourth metatarsal, lying ventromedial-
ly is approximately the same size in all specimens, but is barely distinguishable in D.
bennettianus, and is separated from the larger articular facet for the fourth metatarsal by
a thin tuberous section. In M. rufus however these facets are separated by a small deep
sulcus which channels the medial section of the ligament joining the cuboid to the fourth
metatarsal. The size of the facet on the ventrolateral portion of the cuboid is largest in D.
bennettianus and smallest in M. rufus compared with M. fuliginosus.
The Calcanea of extinct and extant Macropods
Statistical analysis
The results of the cluster analysis can be seen in Fig. 4. This figure is a diagram-
matic representation of the output from the SPSS clustering algorithm. This analysis
included both the fossil specimens and known extant specimens. If any one of the vari-
ables was missing from the measurements of one of the specimens, the respective speci-
men was excluded from the analysis (approximately 15% of the total sample). As a con-
sequence, it was necessary to include the specimens not used in the analysis after the
remainder of the calcanea were sorted out, using these as a guide.
Group “1” contains 14 specimens which are all definitely sthenurine, and most
probably Sthenurus brownei (Metrilees and Porter 1979). There are two distinct size
ranges in this group, and if one species is represented, this may be explained as sexual
dimorphism. (see Fig. 6)
Group “2” contains 12 specimens and is also definitely sthenurine, most probably
Sthenurus occidentalis (Merrilees and Porter 1979). This group contains four very large
specimens, two of which still have their astragalus articulated. There are four medium
sized calcanea and four smaller specimens, suggesting that this group may contain at
least a second species, smaller than S. brownei and S. occidentalis (see Fig.7).
Group “3” contains 14 macropodine specimens. From examination and comparison
with extant specimens, this group contains a majority of specimens attributable to
Macropus fuliginosus, but also interestingly three specimens which can be identified as
Proc. LINN. SOC. N.S.W., 117. 1997
Lo
i)
FUNCTIONAL ANATOMY OF THE MACROPODID PES
20
15
10
= + No rae [es oN on) 00 Valls
Figure 4. Shows the diagramatically simplified results of the cluster analysis performed on the fossilised cal-
canea. Left-hand terminations represent the groups separated in the cluster analysis. This diagram gives a repre-
sentative view of the similarity (indicated by the arbitrary scale of 0-25 on the y- axis) in structure of the
groups clustered out in the analysis
Macropus rufus, an exclusively plains and desert-dwelling kangaroo. Using the measure-
ments taken, the clustering algorithm assigned M. fuliginosus and M. robustus to this
group. Due to the large size of the specimens, this group most likely contains the fos-
silised calcanea of M. fuliginosus rather than the smaller M. robustus.
Group “4” is the largest macropodine group with 57 specimens. Of the extant
material, Macropus parryi, Macropus rufogriseus, and Wallabia bicolor were assigned to
this group through the cluster analysis. M. rufogriseus is one of the best represented
macropodine species identified from cranial elements. There are three distinct size
ranges in this group, the largest of which most probably represents M. rufogriseus.
Group “5” is a large group containing 29 macropodid specimens. Macropus dor-
salis is the only extant species associated with this group through the cluster analysis.
Despite the observation that all of the specimens are roughly equal in size, there are defi-
nitely two distinct morphs in this group, varying in only a few characters which were not
measured in the analysis. Therefore they did not sort or cluster separately.
Group “6” contains six macropodid specimens. Although similar in size to a
female Macropus fuliginosus, no modern macropodid sharing the distinctive features of
these specimens were found. Their form however is somewhat similar to one of the two
morphologically different subgroups of group “5”. Many specimens of this group were
not included in the analysis because of the poor state of preservation common to the
group. Many of the specimens were large yet slender, with many of the processes being
broken or abraded.
Group “7” contains three macropodine specimens. Many of the known extant
species had been clustered in this group; Macropus parma, Macropus irma, Dorcopsus
sp, Petrogale lateralis, Setonix brachyurus, and Macropus eugenii. On comparison with
extant material, only the latter was found to be represented in the fossil collection . It is
interesting to note that so many extant macropodid species are found clustered together
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 33
in one group. This indicates that the genera represented are similar, and that the cluster
analysis was not able to discriminate to the genus level in this cluster.
Group “8” contains 25 macropodid specimens. The majority of this group is proba-
bly attributable to Macropus titan or Macropus giganteus, and also possibly Macropus
robustus. The cluster analysis assigned Macropus fuliginosus to this group. However,
this is a recent lineage, and any such material pre or during the Pleistocene is often
referred to as belonging to Macropus giganteus. There do seem to be two separate
morphs within this group exhibiting very little size variation.
Group “9” is the largest sthenurine group with 34 specimens. It appears that this
group could possibly be divided further into three separate morphs. Based on a compari-
son of the relative abundance of sthenurine cranial elements from the Victoria fossil cave
deposit, the larger part of this group is probably attributable to Sthenurus gilli. This is
supported by the observation that Sthenurus gilli is the smallest species from Victoria
fossil cave deposit, which corresponds with the small size of the specimens. One morph
may be attributed to Sthenurus andersoni (see Fig. 8).
It is now important to understand which factors are causing the main variation of
the morphology of the characters measured. This is achieved through examination of the
principal components analysis.
TABLE 5
The results of the principle components analysis and the first three factors (as defined in text) which account
for the majority of the variation.
Factor Eigenvalue Percent of Variation Cumulative percentage
1 8.926 81.8 81.8
2 0.891 8.1 89.2
3 0.362 33 92.5
Usually with a principal components analysis, a great deal of the variation for the
entire data set can be explained by only two or three factors. Looking at Table 5 we can
see that in this group of calcanea, 92.5% of the variation can be explained by the first
three factors, shown in the cumulative percentage column. The majority of the variation
is undoubtedly in the first factor, which is usually, and certainly in this case, attributed as
size. Hence 81.8% of the variation is accounted for by size.
It can be seen from Fig. 4 that the sthenurine groups cluster separately from the
larger macropodid group. This plot indicates that there were no small sized species
among the sthenurines as seen in groups “4” and “7” of the macropodids. Only principle
components with an eigenvalue greater than ‘1’ are regarded as being statistically signifi-
cant and hence through principal components analysis, size is the only statistically signif-
icant factor.
Comparative morphology of the sthenurine calcanea
As many of the groups sorted out can be assigned to a number of extant taxa, in
which functional and morphological characters are well understood (Owen 1875, Parsons
1896, Windle and Parsons 1897, Craven 1971, Hopwood and Butterfield 1990), I will
concentrate on the sthenurine groups. Where more than one species has clustered into a
single group of the extant macropods, it was possible to discern between the different
species through a visual examination. Within each group there are at least two or more
morphs. These morphs most probably relate to separate species or perhaps separate sexu-
Proc. LINN. SOc. N.S.W., 117. 1997
34 FUNCTIONAL ANATOMY OF THE MACROPODID PES
Figure 5. The right calceaneum of Macropus fuliginosus in dorsal (A), ventral (B), lateral (C), medial (D), and
distal (E) views. DIV- longitudinal division of the medial and lateral facets on the calcaneum; DLF-
Dorsolateral facet; DMF- Dorsomedial facet; FIC- Fibular condyle of the calcaneum; FIF- Facet for the fibula;
GFT- Groove in the sustentaculum tali for the tendon of the Flexor digitorum longus; LFA- Lateral facet for
the astragalus; MFA- Medial facet for the astragalus; PPS- Plantar process of the sustentaculum tali; STT-
Sustentaculum tali; TC- Tuber calcanei; VMF- Ventromedian facet; TPS- Transverse plantar sulcus.
al morphs of the same species. Marked sexual dimorphism is exhibited in modern kanga-
roos by way of size (Poole et.al 1984). As the clustering algorithm used removes the fac-
tor of size, it is expected that sexually dimorphic species will group into the same cluster.
Within group (1) there are at least two morphs; three morphs in group (2); and three
morphs in group (9) (see Figs 6-8). The figures follow after the text.
The tuber calcanei of Macropus fuliginosus is of sub triangular form. Of the
sthenurines, morph (111) of group (9) possesses an ovoid cross sectional shape of the tuber
calcanei (see Figs 5 and 8). The remainder of the sthenurines possess a generally much
broader tuber calcanei which is sub triangular in morphs (1) and (11) of group (1), morph
(1) of group (2), and morph (11) of group (9), compared with the square cross sectional
shape of morphs (11) and (ii1) of group (2), and morph (1) of group (9) (see Figs 6-8). All
of the specimens narrow dorsally to some extent and are flattened plantarly. The tuber
calcanei is flared greatly posteriorly in the sthenurines compared with M. fuliginosus.
The medial and lateral surfaces of the tuber calcanei apex in a longitudinal ridge which is
narrow in morphs (ii) and (iii) of group (9) compared with M. fuliginosus (see Figs 5 and
8). Of the other sthenurines, this longitudinal ridge is moderately narrow in group (1),
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 35
Figure 6. The right calcaneum of a representative of group “1” in dorsal (A), ventral (B), lateral (C), medial
(D), and distal (E) views. APT- Accessory plantar tubercle of the calcaneum; DLF- Dorsolateral facet; DMF-
Dorsomedial facet; FIC- Fibular condyle of the calcaneum; FIF- Facet for the fibula; GFT- Groove in the sus-
tentaculum tali for the tendon of the Flexor digitorum longus; LFA- Lateral facet for the astragalus; MFA-
Medial facet for the astragalus; PPS- Planter process of the sustentaculum tali; STT- Sustentaculum tali; TC-
Tuber calcanei; VMF- Ventromedian facet; TPS- Transverse plantar sulcus.
broad in all morphs of group (2), but most broad and nearly continuous with the medial
and lateral surfaces in morph (1) of group (9). The tuber calcanei is also deepest in morph
(i11) of group (9) relative to its width (see Fig. 8). The tuber calcanei is also shorter in the
sthenurines than M. fuliginosus with the exception of all in group (1) and morph (iii) of
group (9) (see Figs 5, 6 and 8). The sustentaculum tali is relatively narrow transversely in
the sthenurines compared with M. fuliginosus.
A major difference in the sustentaculum tali in dorsal aspect is that in all of the
sthenurines there is a posterior deflection of the plantar portion of the sustentaculum tali
compared with M. fuliginosus, where the plantar portion of the sustentaculum tali is less
angled and deflected medially (see Fig. 5). The sustentaculum tali is very much deeper
Proc. LINN. Soc. N.S.W., 117. 1997
36 FUNCTIONAL ANATOMY OF THE MACROPODID PES
Figure 7. The right calcaneum of a representative of group “2” in dorsal (A), ventral (B), lateral (C), medial
(D), and distal (E) views. APT- Accessory plantar tubercle of the calcaneum; DLF- Dorsolateral facet; DMF-
Dorsomedial facet; FIC- Fibular condyle of the calcaneum; FIF- Facet for the fibula; GFT- Groove in the sus-
tentaculum tali for the tendon of the Flexor digitorum longus; LFA- Lateral facet for the astragalus; MFA-
Medial facet for the astragalus; PPS- Planter process of the sustentaculum tali; STT- Sustentaculum tali; TC-
Tuber calcanei; VMF- Ventromedian facet; TPS- Transverse plantar sulcus.
dorsoplantarly in all of the sthenurine species than in the two macropodine specimens
(see Figs 5-8). Another feature common to all of the sthenurines is that the plantar sur-
face of the sustentaculum tali points directly plantarly compared with anteroplantarly in
group (6) and M. fuliginosus (see Figs 5—8). The plantar process of the sustentaculum tali
makes up part of the plantar surface in morphs (1) and (11) of group (2) as well as morphs
(1) and (ii) of group (9) (see Figs 7 and 8). In the remainder of the sthenurine groups the
plantar process of the sustentaculum tali is separated from the plantar process by a dorso-
plantarly deep groove for the tendon of the flexor digitorum longus. This separation from
the plantar surface is most extreme in morph (iii) of group (9). Another main distinction
between the sthenurines and the macropodines can be seen in the medial profile of the
sustentaculum tali, where in the sthenurines, because of the plantarly deflected plantar
process, a right angle or 90° profile is produced. This is in contrast to M. fuliginosus,
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 37
Figure 8. The right calcaneum of a representative of group “9” in dorsal (A), ventral (B), lateral (C), medial
(D), and distal (E) views. APT- Accessory plantar tubercle of the calcaneum; DLF- Dorsolateral facet; DMF-
Dorsomedial facet; FIC- Fibular condyle of the calcaneum; FIF- Facet for the fibula; GFT- Groove in the sus-
tentaculum tali for the tendon of the Flexor digitorum longus; LFA- Lateral facet for the astragalus; MFA-
Medial facet for the astragalus; PPS- Planter process of the sustentaculum tali; STT- Sustentaculum tali; TC-
Tuber calcanei; VMF- Ventromedian facet; TPS- Transverse plantar sulcus.
where the border is rounded posteriorly and not so steeply inclined, as also noted by
Murray (1995) (see Figs 5-8).
Laterally the fibular condyle is not as wide in sthenurines as it is in M. fuliginosus.
Dorsally, the fibular condyle is substantially variable between the different morphs of the
different groups, but generally the origin of the condyle being more abrupt in the
sthenurines than macropodines. Laterally the fibular condyle bears two scars; one poste-
riorly for the posterior calcaneofibular ligament and one anteriorly for the anterior calca-
neofibular ligament. In morph (i) of group (1) the scar for this ligament encroaches dor-
sally on the condyle, also providing an area for the posterior talofibular ligament which
intervenes in between the fibula and its condyle (Fig. 6). The angle of the scar for the
posterior calcaneofibular ligament is also slightly vertically inclined, but not as much as
in M. fuliginosus (Fig. 5). The posterior border of the fibular condyle is also more abrupt
Proc. LINN. Soc. N.S.W., 117. 1997
38 FUNCTIONAL ANATOMY OF THE MACROPODID PES
in morph (1) than morph (ii) of group (1), compared with M. fuliginosus where the fibular
condyle is curved convexly plantarly and then recurved, demarcating the scar for the
attachment of the anterior calcaneofibular ligament. The fibular condyle is also generally
deeper in the sthenurines than in M. fuliginosus. In all of the sthenurines there is also a
greater area for insertion of the lateral calcaneocuboid ligament.
Dorsally, there are two facets on the calcaneum for articulation with the astragalus.
The origin of the lateral facet for the calcaneum on the dorsal surface is markedly vari-
able in the different specimens. In morph (1) of group (1), the lateral facet for the astra-
galus originates from and is continuous with the dorsolateral surface as in morph (iii) of
group (9), compared with M. fuliginosus, where although the lateral facet for the astra-
galus originates from the dorsolateral surface of the tuber calcanei, it is interrupted by the
higher convex form of the facet. In the remainder of the sthenurines the origin of the lat-
eral facet for the astragalus is dorsally from the tuber calcanei, and is interrupted by its
convex form in morph (ii) of group (1), morphs (1) and (ii) of group (2), and morph (i) of
group (9), compared with morph (i) of group (1), morph (iii) of group (2), and morph (ii)
of group (9) where the origin of the facet is continuous with the dorsal surface of the
tuber calcanei. Immediately anterior to the lateral facet for the astragalus is the fossa for
the proximal ventrolateral process of the astragalus, which is generally much deeper in
the sthenurines. This fossa is also expanded anteroposteriorly in morph (ii) of group (1),
and morph (1) of group (2), rather than being transversely broad in the other sthenurines,
compared with the shallow circular pit in M. fuliginosus (see Fig. 5). In all of the
sthenurines, the lateral facet for the astragalus sits lower than the medial facet. The medi-
al facet for the astragalus is similarly variable between the sthenurines and macrop-
odines, as well as within the sthenurines. Generally the medial facet for the astragalus is
steeper in the sthenurines than in M. fuliginosus, which is shallowly concave on its ante-
rior face and inclined posterodorsally (see Figs 5-8). The lateral and medial facets for the
astragalus are separated by a transversely convex ridge which is particularly high and
narrow in morph (ii) of group (1), morph (1) of group (2), and morph (i) of group (9), and
slightly more shallow but still convex in morph (i) of group (1), morphs (ii) and (iii) of
group (2), and morphs (ii) and (iii) of group (9). This feature may allow a slight degree
of pronation. However in M. fuliginosus the lateral facet sits as high as the medial facet
and is separated from the latter by a small longitudinal ridge, restricting pronation and
supination. Directly behind the medial ridge which articulates with the astragalus, there
is a scar for the posterior talocalcaneal ligament, the main posterior ligament binding the
astragalus to the calcaneum. This scar in sthenurines suggests that the ligament, which is
deflected dorsally and posterolaterally in M. fuliginosus, is deflected only dorsally,
restricting any motion of the astragalus rolling forward. The pit for this ligament is great-
est in morph (ii) of group (1) and morph (i) of group (9) (see Figs 5-8).
On the posterodorsomedial region of the sustentaculum tali is a large scar for the
posteromedial talocalcaneal ligament in the sthenurine groups, perhaps with the exception
of morphs (ii) and (iii) of group (9) where the scar is smaller compared with M. fuligi-
nosus. The sustentaculum tali bears much of the weight of the astragalus in the majority of
sthenurine morphs, with the exception of (ii) and (111) of group (9). It is elongated antero-
posteriorly at its dorsal margin compared with M. fuliginosus where it narrows dorsally.
In dorsal aspect it is also possible to see the varying morphology of the dorsolateral
facet on the calcaneum for the cuboid. In all of the sthenurine groups, the dorsolateral
facet is more narrow transversely, generally shorter anteroposteriorly, but more impor-
tantly the medial side of this portion is deflected laterally, i.e. the calcaneocuboid step is
not as steep as in M. fuliginosus (see Figs 5—8). The dorsolateral facet is shortest in
morph (iii) of group(2) and morph (11) of group (9), which exhibit the shallowest step.
This process is steepest and most elongate in morphs (i) and (iii) of group (9) (Fig. 8).
The remainder of the sthenurines are somewhat intermediate in this character. The shal-
lower the angle of the calcaneocuboid step on the calcaneum, the greater the ability for
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 39
supination of the foot at the calcaneocuboid joint.
Ventrally, the plantar surface of the calcaneum is very greatly elongated in M.
fuliginosus. This is due in great part to the width and position of the transverse plantar
sulcus. This is narrowest and situated anterior most among the sthenurines in morphs (1)
and (ii) of group(1), and morph (i) of group (9), and hence the greatest plantar surface
length. It is also broadest anteriorly in the formerly mentioned morphs and M. fuliginosus
compared with morphs (i) (11), and (ii1) of group (2) and morphs (ii) and (iii) of group
(9). Here the plantar surface is narrow anteriorly and emanates from the lateral side. It is
expanded posteriorly, particularly in morphs (1) and (ii1) of group (2) and morph (11) of
group (9) and results in the plantar surface being nearly triangular in form (see Figs 7
and 8). In M. fuliginosus the plantar surface is nearly uniform in transverse breadth along
its length, expanding only slightly posteriorly. The plantar surface is most narrow in
morph (ii) of group (9). A large component of variation can also be seen plantarly in the
comparative widths of the groove for the tendon of the flexor digitorum longus which is
greatest in morph (11) of group (1), morph (i) of group(2), and morph (1) of group (9) (see
Figs 6, 7 and 8). Correspondingly the groove in the aforementioned specimens is only
shallowly concave transversely compared with M. fuliginosus. Conversely it is deeply
excavated and concave in morphs (11) and (ii1) of group (2), morphs (11) and (111) of group
(9), and to a lesser extent morph (i) of group(1). In all of the sthenurines there is a
demarcation at the base of the sustentaculum tali in the form of a narrow longitudinal
sulcus which is not present in M. fuliginosus.
Distally, there are three facets on the calcaneum which articulate with the cuboid. In
general, the dorsomedial and dorsolateral facets on the calcaneum are more narrow trans-
versely, although still quite broad in morph (i) of group(1) and morphs (11) and (iii) of
group(2), compared with M. fuliginosus (see Figs 5-8). The medial border of the dorso-
medial facet on the calcaneum for the cuboid is steeper in the sthenurine groups and near-
ly vertical in morph (i11) of group (2) compared with M. fuliginosus, which is deflected
dorsomedially. The dorsolateral facet on the calcaneum for the cuboid is also more elon-
gated dorsoplantarly in the sthenurine groups than M. fuliginosus. Also characteristic of
the sthenurines this facet sits lower, or starts more plantarly in distal aspect to the larger
dorsomedial facet, compared with M. fuliginosus where the dorsolateral facet for the
cuboid is equal in height to the dorsomedial facet for the cuboid (see Figs 5-8). The dor-
solateral facet is also deflected dorsomedially in morph (ii) of group (1), morph (i) and
(111) of group (2) and morph (i) of group (9). Where the dorsolateral facet for the cuboid is
expanded in the sthenurine groups, as in morph (i1) of group(1), morph (1) of group (2)
and morph (1) of group (9), it is done so more plantarly, approximately half the way down
the distal surface. The lateral border of the lateral facet for the cuboid is curved convexly
dorsoplantarly in the dorsal region then recurves concavely more plantarly, most strongly
convex dorsally in group (6), morph (ii) of group (1), morph (i) of group (2), and morph
(1) of group (9), and most concave plantarly in morphs (i) and (iii) of group (2), and
morphs (ii) and (111) of group (9) (see Figs 5—8). The dorsolateral facet narrows plantarly
and merges into the ventromedian facet which is generally larger in the sthenurine groups
than M. fuliginosus. In morphs (i) and (11) of group (1), morphs (i) and (ii) of group (2),
morph (i) of group (9), the plantar portion of this facet contributes to the plantar surface of
the pes. In morphs (i) and (ii) of group (2) and morph (1) of group (9), the apex of the ven-
tromedian facet is situated directly underneath the dorsomedial facet compared with the
more medial placement of the facet underneath the step separating the dorsomedial and
dorsolateral facets in M. fuliginosus, morph (iii) of group(2), and morphs (ii) and (iii) of
group (9). It is common to all of the aforementioned morphs that the ventromedian facet
plays no direct weight bearing role. Common to all of the sthenurine groups is the curved
form of the ventromedian facet, whose dorsal and plantar borders are curved transversely
such that the facet is “u” shaped in morphology compared with M. fuliginosus, which pos-
sesses a small ovoid facet. Also common to the sthenurine groups is the merging of the
Proc. LINN. Soc. N.S.W., 117. 1997
40 FUNCTIONAL ANATOMY OF THE MACROPODID PES
ventromedian facet on to the dorsomedial facet for the cuboid, making the three facets
continuous, and forming a fossa in the centre of the three facets, compared with M. fuligi-
nosus Where the two facets are separated by a deep groove (see Figs 5-8).
DISCUSSION
The plesiomorphic characters of the marsupial pes
In constructing the cladistic relations of this group, it was revealed that there were
no phylogenetic reversals in the cladogram. The pes of the outgroup expressed fourteen
of the seventeen characters that were considered to be symplesiomorphic. This indicated
that the morphology of the pes is highly conservative. There were only three characters
where the polarity could not be determined. These were: | - (Character 10) The tuber
calcanei, which was phylogenetically uninformative due to its diverse morphology.
Murray (1995) similarly found this character to be uninformative in his study of the phy-
logenetic relationships of the late Miocene kangaroo, Hadronomas. 2 - (Character 5) The
possession of a trochlear notch along with the (plesiomorphic) long astragalar head and
neck in the wombat can be attributed to parallelism of evolution. Similarly the conver-
gence in form of the sustentaculum tali of the koala and Hypsiprymnodon is unlikely to
be the result of similar function in these species. The expression of this character state in
the koala may be attributed to the retention of an ancestral character state. 3 - (Character
11) Based on a single feature, the Potoroidae would appear to be derived from the
Phascolarctidae, contradictory to current phylogenetic placement. The contradictory
nature of the cladogram can be explained by the use of a limited character set, i.e. the
groupings of the outgroup marsupials are being determined on only one or two characters
which are seen to be divergent from the generalised marsupial pes. This is most probably
biased through the choice of characters which seemed to be most divergent in the
macropodid species. A larger set of characters would have to be used for a more compre-
hensive study, particularly for the plesiomorphic states.
There were six shared derived features which unite Hypsiprymnodon (the most
Plesiomorphic of the living kangaroos) with M. fuliginosus. This is consistent with the
current placement of this family and its contained species, as intermediate in structure
between the phalangerids (possums) and the macropodids (kangaroos). Disregarding the
uninformative characters (5,10 and 11), there were no reversals of characters between the
ancestral marsupial grade and that of M. fuliginosus.
The derived features of the macropodine pes are considered to be important, as
they define its specific structure in relation to the generalised marsupial pes. Those fea-
tures which are characteristic of the macropodines are likely to be related to locomotion
and habitat preference. A more complete analysis of the features of the generalised
macropodine form was attained through an examination of Macropus fuliginosus which
is considered to be derived relative to Hypsiprymnodon. It is necessary to understand the
functional nature of unique derived features of the macropodine pes.
Anatomy of the Hindlimb and Pes of Macropus fuliginosus
The results of this work support earlier findings (Craven 1971, Parsons 1896
Adnams-Hodges 1988, Hopwood and Butterfield 1990) that the main contribution to the
muscle mass in the macropodines is in the flexor and extensor muscles of the hindlimb.
This reflects the specialised form of locomotion. The actions of the separate muscles of
the crus on the pes have been further described. The insertion points of the tendons and
ligaments produce scars or rugosities on the surface of the bones. This knowledge has
been used to identify homologous features on the bones of the extinct forms.
Proc. LINN. SOc. N.S.W., 117. 1997
N. BISHOP 4]
The functional anatomy of the western grey kangaroo, Macropus fuliginosus was
used in this study as a general model against which to compare other macropodid species.
Also studied was the anatomy of the binding ligaments of the pes, an area which
seems to have been ignored in previous studies. The main muscles acting on the pes have
lines of action predominantly in the sagittal plane. Ligaments of the pes also tend to
restrict motion to the sagittal plane. Ligaments also restrict the elongated pes from being
excessively dorsiflexed. These ligaments are the large posterior and anterior calcaneofibu-
lar ligaments in lateral aspect, and the posterior tibiotalar ligament and tibiocalcaneal liga-
ment in medial aspect. Their cruciate (crossed) form stabilises the ankle. Similar results
were found by Lewis (1983), and by Parsons (1896) who highlighted the stabilising nature
of the “X” formation in the rock wallaby, Petrogale xanthopus. Parsons (1896) also iden-
tified two calcaneocuboid ligaments, a large plantar ligament, referred to as the ‘outer lig-
ament’ running from the rugose plantar surface of the calcaneum to the lateral tubercle of
the cuboid, and on to the base of the fourth and fifth ligaments; ‘the medial ’, being small-
er but still prominent, passing from the sustentaculum tali to the cuboid and the base of
the second and third metatarsals. These results complement the findings of this research. I
conclude that the function of these ligaments is to prevent over dorsiflexion of the pes,
and allow for the possibility of elastic recoil, loaded when the animal lands.
The most complete description of the macropodine foot to date is by Owen (1875)
in his description of the pes of Macropus rufus. The work of Owen (1877-1878) supports
the current findings, in that the narrow form of the pes is related to its saltatory mode of
locomotion. Lewis (1980, 1982) also notes the form of the astragalus (a functionally
important bone of the ankle) as being remodelled into a trochlear shape, convergent in
function with the placentals, supporting the current findings. Conclusions regarding the
splint-like nature of the fibula in macropods and its role in restricting the motion about the
ankle to the sagittal plane are supported by Barnett and Napier (1953). One peculiarity of
this form in macropods is that while the trochlear ridges of the astragalus are parallel, they
are slightly oblique to the long axis of the foot, resulting in internal rotation of the foot.
This may be linked to the idea expressed by Barnett and Napier (1953), Parsons (1896),
and Wells and Tedford (1995), of the possibility of a spring return of the fibula when
loaded as the lateral condyle of the femur forces the head posteriorly during flexion of the
knee. With the pes firmly placed on the ground, the tibia, fibula and remainder of the ani-
mal passes over the fixed pes, and the angle of the trochlear crests will act to rotate the
crus externally, such that the action of the femur on the fibula is magnified. Parsons
(1896), Windle and Parsons (1897), and Hopwood and Butterfield (1990) also provide
evidence on the general morphology of the macropodine pes which supports the current
study. The transverse nature of the subtalar joint complex was identified supporting earlier
findings of Owen (1877-1878), Lewis (1964, 1980, 1983), and Barnett (1970).
Modifications of the macropodine pes.
Despite being restricted by the specialised mode of locomotion and necessary
structural modifications associated with it, the modern macropods have managed to
occupy a number of diverse habitats; e.g. as Macropus rufus, a plains dwelling kangaroo,
and Dendrolagus bennettianus, a kangaroo adapted for an arboreal lifestyle.
Modifications of the Macropodid Pes for Open Plains and Arboreal Habitats
The main difference of the pes of Macropus rufus, an open plains animal, from that
of Macropus fuliginosus, is the gracile form of the entire pes. As M. rufus inhabits large
areas of flat treeless rolling plains of calcareous clay soils it is often referred to as the
“open plains kangaroo”. M. rufus also has the highest recorded speeds among modern
macropodines (Bennett 1987).
There are a number of features of the pes of Dendrolagus bennettianus which I con-
Proc. LINN. Soc. N.S.W., 117. 1997
42 FUNCTIONAL ANATOMY OF THE MACROPODID PES
sider to be adaptations to an arboreal lifestyle. Despite the macropodines having evolved
from the arboreal possums, there is little doubt that Dendrolagus is a macropodine and not
simply an ancestral grade between the possums and the kangaroos. It has clearly secon-
darily modified a terrestrial hindlimb for an arboreal lifestyle. Those features considered
to be derived in the macropodine pes are also expressed in D. bennettianus.
i) Modifications of the Astragalus: The main distinction between Dendrolagus and
Macropus fuliginosus is the form of the transversely broad astragalus. This feature is in
accordance with the general breadth of the entire pes of this genus. The broad pes is a
functional requirement for the arboreal habitat, giving greater stability on narrow tree
limbs. The parallel trochlear crests are oblique to the long axis of the foot. In this respect
the form is convergent in function with the other arboreal species examined in this study,
the possum and koala. The oblique nature of the articulation of the crus on the pes results
in one of two actions. With the proximal portions of the hindlimb held close to the body,
when climbing, the oblique trochlear ridges cause the foot to be internally rotated, con-
ceivably aiding in climbing tree trunks. Similarly when the feet are closely placed next to
each other, such as when sitting on a branch, the knees are externally rotated, lowering
the body and centre of gravity, resulting in a more stable posture.
Conforming with the remainder of the pes, in M. rufus the astragalus is relatively
narrow transversely, and elongated anteroposteriorly. It is probable that the great antero-
posterior length of the astragalus and of the trochlear ridges have the effect of restricting
motion of the pes to the sagittal plane. The parallel trochlear ridges are in line with the
longitudinal axis of the foot, indicating that any movement of the pes is confined to the
sagittal plane. Another feature of this articulation in M. rufus is the medial trochlear
ridge, which is greater in height than the lateral trochlear ridge and associated with a well
demarcated and steep medial malleolus. This feature seems to be related to maintaining
the pes in the sagittal plane during locomotion. Another factor restricting motion at this
joint is the line of action of the main medial ligaments. Medially, the insertion of the two
crossed ligaments on the astragalus are defined by a long horizontal ridge. This arrange-
ment limits external and internal rotation of the pes as well as pronation.
The trochlear groove for the tibia is deepest in Dendrolagus, which I believe is due
to the necessity for greater mobility at the ankle, in particular in being able to pronate
and supinate the foot. When viewed in distal aspect, the medial and lateral trochlear
ridges are equal in height compared with Macropus fuliginosus where the medial
trochlear ridge is higher. I believe this is also related to the ability to pronate and supinate
the foot. Barnett and Napier (1952), Hicks (1953), Close (1956) and Sarrafian (1993)
have examined the nature of the ankle articulations in human specimens. Their results
support the findings of this work that flexion and extension of the foot also affects other
motions, such as pronation, and supination. Barnett (1970) and Lewis (1980) note the
transverse nature of the subtalar joint complex, and Barnett (1970) notes the diverse mor-
phology of this feature among the modern macropodines, but makes no functional inter-
pretations.
Ventrally the articular facets of the calcaneoastragalar articulation are transversely
broad, and this is most likely related to the general breadth of the pes. In Dendrolagus
the medial malleolar tibial articulation is ovoid, expanded anteroposteriozly, and is prob-
ably related to greater mobility of the tibia. In Dendrolagus the scar for the posterior
tibiotalar ligament suggests that the action of this ligament would most likely allow for
greater eversion and inversion of the foot. Similarly the area for attachment for the ante-
rior tibiotalar ligament is much reduced, possibly allowing for greater supination when
the foot is plantarflexed, although even the reduced ligament would constrain this range
of motion. The navicular facet on the astragalus is transversely wide, corresponding with
an enlarged fifth digit. Lewis (1980) notes the nature of an intra articular meniscus in
possums, which is much reduced in macropodids. Lewis (1980) also suggests the menis-
cus provides greater mobility at the articulation between the fibula, tibia, and the astra-
Proc. LINN. Soc. N.S.W., 117. 1997
N. BISHOP 43
galus. Examination of a dried ligament preparation from the South Australian Museum
indicated that the meniscus is enlarged in Dendrolagus compared to other macropodines.
11) Modifications of the Calcaneum: The general morphology of the calcaneum of
Dendrolagus corresponds with the form of the rest of the pes, being broad and dorso-
plantarly compressed. The tuber calcanei is ovoid, reminiscent of the form of the pos-
sum, and considering its similar habitat, may also be convergent in function. Similarly,
the sustentaculum tali is extremely broad, tapering medially as in the koala and possum,
also related to the breadth of the pes. The sustentaculum tali bears a groove for the ten-
don of the flexor digitorum longus, the function of which is to flex the digits. The mor-
phology of this groove in Dendrolagus 1s such that the tendon would be relatively uncon-
strained, allowing the digits to be flexed in a greater range of positions. There is a
reduced articulation of the fibula with the calcaneum in Dendrolagus, as noted by
Barnett and Napier (1953). This would suggest greater mobility at the articulatio tar-
socruralis (between the crus and the pes) as the constraining nature of the talofibular
articulation is minimised. This is associated with only a slight lateral projection for the
fibula articulation.
In M. rufus the tuber calcanei shows evidence of being adapted to a biomechanical
function, associated with an open plains habitat, as it narrows dorsally. Ontogenetic
changes of bone are related to the direction and type of forces placed on it (Hildebrand
1988). As the tuber calcanei narrows dorsally, to a shape which is essentially triangular,
it is indicative of the unidirectional forces acting upon the calcaneum. The groove for the
tendon of the flexor digitorum longus runs under the very narrow sustentaculum tali,
suggesting that the digits are only flexed when the foot is in the sagittal plane of the
body, i.e. only when there is a direct line of action for this tendon. Dorsally, the large
area for insertion of the ligamentum cervicis tali is associated with the strong binding of
the astragalus to the calcaneum. This further restricts motion at the subtalar joint.
Another feature of the calcaneum suggesting a restriction of motion to the sagittal
plane, is the size of the articular facet for the fibula. The fibula and the associated liga-
ments ensure the crus is securely braced to the pes, and restricts any internal or external
rotation of the foot about the tibia. There are also a number of features of the ankle of M.
rufus that relate to restriction of lateral movement at the calcaneocuboid articulation.
This is supported by the arrangement of the ligaments at this joint. The dorsolateral facet
of the calcaneum for the cuboid is anteroposteriorly compressed, but more importantly
the step between this and the dorsomedial facet is well defined. This feature constrains
any pronatory or supinatory motions of the pes.
Ventrally, the plantar surface of the pes is particularly tuberous, providing a large sur-
face area for insertion of the plantar calcaneocuboid ligament. Medially, the sustentaculum
tali is also tuberous for the insertion of a large medial calcaneocuboid ligament. These find-
ings concur with those of Parsons (1896). The areas for insertion of the ligaments binding
the cuboid to the fourth and reduced fifth metatarsal are also correspondingly large.
Dorsally, of the two articulating facets for the astragalus, the medial possesses a
ridge dorsally which is straight, runs transversely, and is deflected slightly anteriorly. The
function of this is probably for greater flexion and extension of the pes at the subtalar
ankle joint. Directly behind this ridge is a large scar for the posterior talocalcaneal liga-
ment, suggesting that the astragalus is well bound to the calcaneum posteriorly, resisting
excessive pronation of the pes. A further feature of the talocalcaneocentralis joint of
Dendrolagus is the fossa for the anterior proximal process of the astragalus is not well
developed, and hence motion at this articulation is not restricted.
In summary, the features of the subtalar joint complex of Dendrolagus allow for
greater freedom of movement associated with an arboreal habitat. Distally the main fea-
tures of the calcaneum that allow for increased motion at the distal joint are related to the
shallow gradient of the normally steep facets for the cuboid as expressed in other
macropodines. The basic form of the distal articular surface is shallowly convex, both
Proc. LINN. Soc. N.S.W., 117. 1997
44 FUNCTIONAL ANATOMY OF THE MACROPODID PES
transversely and dorsoplantarly (cup shaped), and corresponds to what Sarrafian (1993)
describes as a male ovid surface. The motions generated at a male and female ovid (or
ball and socket) joint are those of flexion, extension, pronation, supination, internal rota-
tion, and external rotation (Sarrafian 1993).
11) Modifications of the Cuboid: The proximal articulating facet of the cuboid cor-
responds with the calcaneum as the female ovid surface. As with the calcaneum and
astragalus, the cuboid 1s transversely broad and dorsoplantarly compressed. Distally, the
separate articulating facets on the cuboid for the fourth and fifth metatarsals take the
form of a single facet. In medial or lateral view, the cuboid is rhomboidal in appearance,
extended anteriorly dorsally, the function of which would be to prevent excessive dor-
soflexion of the foot, being restricted by the protruding dorsal portion. Ventrally, in
Dendrolagus, the medial plantar crest of the cuboid is enlarged, and in contact with the
substrate surface, providing greater stability for the pes.
In summary, the feature of the Dendrolagus pes provides greater stability in the
width of the pes in addition to greater mobility of the ankle joints, both necessary for an
arboreal habitus.
These results suggest the main adaptations for an open plains habitat are an elon-
gated narrow pes with motion restricted to the sagittal plane. The comparison of these
two forms (arboreal and terrestrial), clearly indicate a large degree of homoplasticity in
the features deemed to be derived within the macropodines. This has allowed the
macropodines to radiate into a number of different habitats.
The Cluster Analysis
One aim of this study was to sort the fossilised calcanea from the Victoria Fossil
Cave, Naracoorte. From an examination of both the cluster analysis and principal com-
ponents analysis it can be seen that sthenurines separate from the macropodines and form
three distinct groups. A closer examination of the group revealed a number of features of
the calcaneum which were species specific. In a similar fashion, it is possible that the
sthenurine groups may include more than one species.
The cluster analysis was an invaluable tool in determining primary structural dif-
ferences in the large number specimens examined. It was not until after a discrete func-
tional and structural analysis had been carried out that a number of “species specific”
characters could be recognised.
Functional Anatomy of the Sthenurine Pes
Very few studies (Stirton 1963, Tedford 1966, 1967; Adnams-Hodges 1988;
Murray 1991, 1995; Wells and Tedford 1995) have attempted to determine the functional
nature of the postcranial elements of the sthenurines. Wells and Tedford (1995) examined
the postcranial remains of three species of extinct sthenurines and suggested that as there
are no modern descendants it was difficult to determine the locomotory habits of these
animals for lack of a reliable functional analogue. These authors were however able to
relate a number of features relating to the locomotor ability of the sthenurines. Wells and
Tedford (1995) suggest that the sthenurine’s mode of locomotion was of limited manoeu-
vrability, and further suggest that adaptations of the hindlimb and pes are related to sup-
porting the weight of the animal, both during locomotion and stand-up browsing.
Just as the modern macropodine forms have diversified into a wide range of habi-
tats, with accompanying differences in foot structure, so it is expected that differences in
morphology of the calcaneum of the extinct forms may also be related to habitat.
The morphology of the tuber calcanei is much varied in the sthenurine morphos-
pecies. Despite being generally more robust than Macropus, the four morphospecies share
a subtriangular form; three of which have a square cross-sectional shape, and the fourth is
Proc. LINN. SOc. N.S.W., 117. 1997
N. BISHOP 45
transversely narrow. The three ‘species’ where the tuber calcanei 1s square 1n cross-section
suggest that forces acting on this process of the calcaneum are both in the sagittal and
transverse planes. The transversely narrow tuber calcanei is indicative of forces being
directed largely in the sagittal plane only. With the exception of four species, the tuber cal-
canei of the sthenurines is relatively anteroposteriorly compressed. This would suggest
that the sthenurines were not as well adapted for fast locomotion as many of the extant
macropodines. Their form is divergent from the form of the highly specialised red kanga-
roo, Macropus rufus. As noted by Tedford (1966, 1967), Adnams-Hodges (1988), Murray
(1995), and Wells and Tedford (1995), the sustentaculum tali is generally narrower in the
sthenurines than the macropodines, with the exception of two morphospecies where the
sustentaculum tali is particularly broad posteriorly. This seems to be convergent in func-
tion with the possums and koalas, in being an adaptation allowing the digits to be plantar
flexed while the foot is internally rotated. This is supported in other features of the susten-
taculum tali. In medial aspect, the profile of the sustentaculum tali is dorsoplantarly deep
and right-angled, providing two possible functions. While the main extensor muscles, the
gastrocnemius and flexor digitorum brevis provide the majority of the power to extend the
pes during standing, the flexor digitorum longus may augment this action. As the animal
starts to stand, the distal digits are firmly planted on the ground, and the action of the ten-
don of the flexor digitorum longus running underneath the pes would be to elevate the pes
at the point of the sustentaculum tali. The right angled form of the sustentaculum tali
would also impede the tendon from dislodging medially. This is supported by another fea-
ture of the sustentaculum tali in that it is pointed plantarly, further constraining the tendon
with the foot extended, and also possibly providing support medially, as it contributes to
the plantar surface of the pes in all but one morphospecies.
Also related to the action of the flexor digitorum longus tendon across the sus-
tentaculum tali is the presence of a small longitudinal groove lateral to the path of the
tendon, separating the sustentaculum from the body of the calcaneum plantarly, also
reported by Wells and Tedford (1995). Deep within this groove is a small foramen,
which suggests that this adaptation is for protection of the blood and nervous innerva-
tion of the calcaneum. Another feature of the calcaneum related to raising the body is
the degree to which the tuber calcanei is flared posteriorly. On comparison of
sthenurine and macropodine specimens of equal size, this feature indicates large areas
for insertion of the tendons of the gastrocnemius and flexor digitorum brevis, suggest-
ing that the action of these tendons in extending the pes is particularly important in the
sthenurines, supporting earlier findings (Adnams-Hodges 1988, Murray 1989, 1995,
and Wells and Tedford 1995).
The fibular condyle is reduced in the sthenurines and relatively flat compared with
M. fuliginosus, indicating less fibular contact with the calcaneum. On many of the
sthenurine calcanea, the fibular condyle is depressed, sitting lower on the lateral face.
This may indicate that the foot is able to be pronated to a greater degree, supporting the
suggestion of Murray (1995) that the majority of the weight is transferred to the medial
side in the sthenurines. This is further supported by a number of features of the calca-
neum. If the sthenurines were to distribute the majority of their weight medially, with
their foot pronated, we would expect to see a number of mechanisms preventing the
astragalus from being displaced from the calcaneum. One of the most notable pieces of
evidence for this is the large scar for the calcaneoastragalar ligament, which is similarly
large in the arboreal forms, possum, koala, and tree kangaroo. This was also noted by
Stirton (1963), and Murray (1995). This ligament restricts anteromedial movement of the
astragalus about the calcaneum. The medial articulation for the astragalus is greatly
accentuated and bears a deep scar for the posteromedial talocalcaneal ligament, further
limiting medial displacement of the astragalus. There is a ridge between the medial and
lateral facets for the astragalus, which is very broad in two of the morphospecies. This
ridge also forms the medial border of an extremely deep pit which is anteroposteriorly
Proc. LINN. Soc. N.S.W., 117. 1997
46 FUNCTIONAL ANATOMY OF THE MACROPODID PES
elongated in some species, bearing the anterior proximal process of the astragalus. This
feature is similarly related to stabilising the subtalar ankle joint, imperative not only in
locomotion, but also to support the huge bulk of the body when standing erect. This pro-
vides an explanation for the reduction of the fifth metatarsal as the majority of the weight
is being borne on the medial side of the pes.
The fifth metatarsal is expanded proximally (Adnams-Hodges 1988; Murray 1991,
1995; Wells and Tedford 1995), and may relate to morphological differences of the distal
lateral portions of the calcaneum. The entire distolateral portion in the calcaneum is
depressed in distal aspect, as well as being angled anteroplantarly, transmitting weight
through to the expanded proximal portion of the fifth metatarsal. In four of the sthenurine
morphospecies, there is a large transverse plantar sulcus on the plantar surface of the cal-
caneum, convergent in form with the potoroids.
The tuberous section of the plantar surface provides insertion for the plantar calca-
neocuboid ligaments, which tightly bind the calcaneum to the base of the cuboid, fourth
and fifth metatarsal. The sthenurine’s possession of an elongate transverse plantar sulcus
may be related to utilisation of the elastic properties of the ligament originating posteri-
orly on the calcaneum. As the ligament is not bound to the calcaneum at the plantar sul-
cus, the ligament may act like a bow across the plantar surface between the calcaneum
and the cuboid.
In the sthenurines as with Macropus rufus there is evidence for large lateral and
medial calcaneocuboid ligaments, suggesting reduced lateral movement. Mobility at the
calcaneocuboid articulation is also heavily influenced by the morphology of the distal
articulating facets for the cuboid. In two of the morphospecies the dorsolateral facet for
the cuboid is particularly narrow, but also broad in three of the morphospecies. The most
important functional nature of the calcaneocuboidal step, which is shallow in four of the
morphs, suggests the foot may be supinated or pronated at the calcaneocuboid articula-
tion. The dorsolateral facet for the cuboid is also anteroplantarly compressed in a few of
the sthenurine groups, convergent in form with M. rufus, the distinction being in the
degree of truncation of the step. A further consistent feature of the sthenurines is that the
dorsomedial and ventromedian facets are fused, surrounding a circular fossa suggesting
increased rotational mobility at the calcaneocuboid articulation. The ventromedian facet
for the cuboid is much larger in all of the sthenurines than the macropodines. This would
be attributed to its weight bearing role during stand-up browsing. In three of the mor-
phospecies, the plantar apex of the ventromedian facet is situated directly below the dor-
somedial facet, compared with a more lateral placement of the ventromedian facet,
below the calcaneocuboidal step in the other morphospecies. The placement of this fossa
indicates where the weight is being borne in the foot, and that, in the former group, the
foot was stressed in a slightly pronated position.
Inferred Locomotory and Feeding Habits of the Sthenurines from the Naracoorte Area
On discrete analysis of the sthenurines calcanea, at least eight morphospecies could
be identified. The diverse morphology of the calcaneum suggests that they were segre-
gated into a wide range of habitats.
Inferred Habits of Group (1):
There are two morphospecies within this group which were originally thought to
represent two sexual morphs of Sthenurus browneii (Merilees and Porter 1979). It was
not within the scope of this study to include a complete comparison of the sexual dimor-
phism present in the modern macropodine species, which should be carried out to test
such a hypothesis.
It can be inferred from the pes structure that this sthenurine group is adapted main-
ly for bearing the large weight of the animal during locomotion. The sthenurines in this
Proc. LINN. SOC. N.S.W., 117. 1997
N. BISHOP 47
group were not able to rotate their feet to the extent of the modern macropodines.
Because of the monodactyl nature of the pes, it is concluded that these sthenurines were
adapted to a habitat relatively free of obstacles. There are still adaptations apparent to
align the foot under the body in the sagittal plane during locomotion.
Inferred Habits of Group (2):
There are three morphospecies in this group. Because the measurements were not
able to discriminate to species level, morph | of this group has the same relative propor-
tions as the other two. On further examination morph | was found to be anatomically
very different. There are a number of features discriminating the second two morphs of
group (2) which are probably Sthenurus occidentalis based on a comparison with a
known specimen from the Green-Water Hole at Tantanoola, S.A.:
It can be inferred from the morphology that the pes of this sthenurine group is
adapted for bearing the large weight of the animal (approximately 20% greater in mass
compared with the largest of the modern macropodine species) during locomotion, and
that there are adaptations for greater mobility at the ankle. It would seem that the
sthenurines in this group have compensated for the loss of the stabilising lateral digits by
having greater ability to control the motion of the foot. The loss of stability at the ankle
however would indicate slow locomotion as much of the action of the muscles would be
in control of the foot, particularly through uneven terrain. These sthenurines also would
have been well adapted for stand-up browsing.
Morph | of group (2) shows many adaptations to keeping the pes in line under the
body during locomotion, but also show adaptations for being able to rotate the foot inter-
nally and externally. These features are most likely related to control of the foot during
stand-up browsing, such as moving the digits and the feet, particularly with the feet exter-
nally rotated, providing greater stability. The sthenurines of this morphospecies were prob-
ably also moderately well adapted to locomotion through habitats with uneven terrain.
Inferred Habits of group (9):
The detailed anatomy of the morphs of group (9) is extremely varied. morph (i) is
concluded to be a sexual morph of group (1) morph (ii) with the exception of the com-
paratively short tuber calcanei. Morph (ii) of group (9), which is surmised to be
Sthenurus gilli, can be distinguished by the following characters:
The features of morph (ii) of group (9) (tentatively Sthenurus gilli) suggest that
this species was particularly well adapted to stand-up browsing, being able to externally
rotate the foot, producing a stable posture. This species also possessed features allowing
the foot to be pronated and supinated. This species had stable upper and lower joint com-
plexes, suggesting that during locomotion the feet were able to be placed underneath the
body, possibly through internal rotation of the crus about the articulation with the femur.
It is evident that this smaller species does not possess the weight bearing adaptations.
This is to be expected considering the small size of this species.
Morph (iii) of group (9) diverges significantly from the general robust form of the
remaining sthenurines.
It is evident that this species bears few of the adaptations to locomotion and stand-
up browsing recognised in the remainder of the sthenurines. It is proposed that this
sthenurine species was particularly gracile and had proficient locomotor capabilities. The
depth of the tuber calcanei suggests that the animal still had to bear a substantial weight
load, which was subsequently directed primarily in the sagittal plane. There was limited
mobility of the foot, extraneous to motion in the sagittal plane. Evidence suggests that
during locomotion, the feet did not have to be forced underneath the body to support a
great weight. It is concluded that this species was adapted to a plains environment, and
was the most gracile or slender of the sthenurines. This sthenurine species was not partic-
ularly well adapted to stand-up browsing.
Proc. LINN. Soc. N.S.W., 117. 1997
48 FUNCTIONAL ANATOMY OF THE MACROPODID PES
CONCLUSIONS
The shared pedal characters of the diprotodont marsupials were compared with
other diprotodont marsupials including the possum, wombat and koala. Using these as an
outgroup, the musky rat-kangaroo, Hypsiprymnodon was shown to possess the fewest
derived characters of the macropodids. The western grey kangaroo, Macropus fuligi-
nosus was shown to be very derived relative to the condition expressed in
Hypsiprymnodon.
A study of the functional anatomy of the hindlimb and pes of Macropus fuliginosus
demonstrated that the derived morphology related directly to its specialised bipedal mode
of locomotion. Some of the macropodines have diverged, and are present in varied habi-
tats. Dendrolagus bennettianus, the tree-dwelling kangaroo, possessed a short broad pes,
and an ankle structure with high mobility, clearly adaptations to an arboreal habitat.
Macropus rufus, the plains-dwelling kangaroo showed adaptations such as a narrow
elongated pes with a highly constrained ankle, which relate to locomotion in the even ter-
rain plains habitats.
A cluster analysis of the fossil calcanea sorted the sthenurines from the macrop-
odines. The sthenurines could further be broken down into nine distinct morphological
groups, some of which could be assigned to particular species.
The main features uniting the sthenurines suggest that the sthenurine pes was
adapted for stand-up browsing. While there is no doubt that like the modern kangaroos,
the sthenurines utilised the bipedal hop for locomotion, the results support earlier find-
ings that their morphology suggests that the feet were adapted to bearing the great weight
of the animal during locomotion.
However, some of the sthenurine calcanea, while still possessing adaptations for
stand-up browsing were noted to be particularly slender, and lacked the adaptations relat-
ed to weight bearing. Evidence suggests that the sthenurines from the Naracoorte area
were partitioned into several different habitats, from scrub and uneven terrain to plains
environments.
The distinct morphological variance exhibited in the characters of the pes of these
sthenurine forms suggest it may be possible to reconstruct the phylogeny of these groups
based on anatomy of the foot.
AKNOWLEDGEMENTS
I wish to thank all of my colleagues at Flinders University who helped me with this study; Assoc. Prof.
Rod Wells, Assoc. Prof. Russel Baudinette, Gavin Prideaux, Matthew McDowell, and Pyramo Marianelli. I
would also like to thank Lynette Queal and Neville Pledge of the South Australian Museum for providing me
with extant and extinct specimens respectively. I would like to thank the Cleland Wildlife Park for provision of
a number of carcases for dissection. I would also like to thank Dr. Michael Schwartz for advice and help with
the statistical analysis.
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Proc. LINN. SOC. N.S.W., 117. 1997
The Late Quaternary Sediments and Fossil
Vertebrate Fauna from Cathedral Cave,
Wellington Caves, New South Wales
LYNDALL DAWSON AND M.L. AUGEE
Biological Science, University of New South Wales, Sydney, NSW 2052
Dawson, L., and Augee, M. L. (1997). The Late Quaternary sediments and fossil vertebrate
fauna from Cathedral Cave, Wellington Caves, New South Wales. Proceedings of the
Linnean Society of New South Wales 117: 51-78
Excavation of the floor of Cathedral Cave, Wellington Caves, was undertaken between
1982 and 1986. Three major phases of deposition are recognised, representing the end of the
last interglacial (Phase 1), the last glacial maximum (Phase 2) and a Holocene phase (Phase 3).
Radiocarbon dating of small amounts of charcoal in Phase | gave dates ranging from 33,800 BP
to 21,000 BP, while dates of 2,590 BP and 2,950 BP were obtained from Phase 3 sediments.
Within these Phases stratigraphic, sedimentological and taphonomic changes are appar-
ent. Phase | has the characteristics of an attritional entrance facies deposit. Large taxa (>1.5 kg
body weight) are represented by juveniles only, except for scavengers and carnivores, suggest-
ing a pitfall trap. The abundance of small mammals is consistent with their accumulation due
to owls and the Ghost Bat, which is present in the bottom 1.25 m of the deposit. In Phase 2 a
high degree of disturbance is suggested by the inclusion of large limestone boulders and rocks
and the highly fragmented nature of the bone. Two distinct heavily indurated ‘floors’ dip
toward the south-east and suggest entry of the sediments from a point in the roof of the cave
co-incident with the apex of the present ‘altar’ formation. The sediments of Phase 3 are not
indurated and contain no large boulders or bone fragments, larger species being represented by
isolated teeth only. The entrance source of Phase 3 sediments is unknown.
At least 38 marsupial taxa have been identified of which 10 are extinct. Of the remain-
ing 28 extant marsupial taxa only 9 may still occur west of the Great Dividing Range today.
Of the non-marsupial taxa rodents are most abundant. Microchiropteran bats are represented
in several strata, as are reptiles and small birds. Pleistocene ‘megafauna’ taxa are represented
in Phase 1 and Phase 2. “Megafaunal’ species include Macropus (Osphranter) altus, M. (M.)
titan, M. (M.) ferragus, M. (Notamacropus) n. sp., Protemnodon sp., Simosthenurus oreas,
and Thylacoleo carnifex.
The fauna of medium and smaller sized mammals exhibits disharmonious assem-
blages typical of other late Pleistocene/Holocene faunas of eastern and south-eastern
Australia. The presence of the Ghost Bat, Macroderma gigas, in Phase | suggests a suitable
configuration of the cave and cave entrance at that time, and a warmer, wetter climate than
Phase 2. Fauna of Phase 3 lacks extinct species and is representative of conditions which
existed at the time of first European settlement of the Wellington region.
Manuscript received 18 June 1996, accepted for publication 23 October 1996.
KEYWORDS: Marsupial, Megafauna, Quaternary, Wellington.
INTRODUCTION
The Wellington Caves, located 6 km south of Wellington on the western slopes of the
Great Dividing Range in New South Wales, have been renown, since their first discovery
in 1830, as a rich source of fossil bones, including many taxa of extinct marsupials. The
history of their exploration and excavation has been reviewed most recently by Dawson
(1985) and Osborne (1991). Descriptions of the caves complex, the associated sediments
and stratigraphy have been published by Frank (1971), Francis (1973) and Osborne (1983).
The complex consists of several natural chambers and man-made tunnels which
have been excavated from the sediments which fill much of the cave system. Over a peri-
od of more than 150 years, fossil bones have been collected from Mitchell Cave,
Proc. LINN. Soc. N.S.W., 117. 1997
52 LATE QUATERNARY SEDIMENTS
Lower
Chamber
Sete
(go age
Water eX
o CEN —
0 Main Chamber
Position of
UNSW
Excavation
Nth.
5 metres
MAIN ENTRANCE mR,
Blocked
Entrance
Figure 1. Map of the floor of Cathedral Cave, Wellington Caves, indicating the position of the University of
New South Wales’ excavation (1982-1986).
Cathedral Cave, Big Sink, Bone Cave and the Phosphate Mines. It had previously been
assumed that all the sediments in the caves were of Pleistocene age because of the pre-
dominance in museum collections of the bones of extinct marsupial megafaunal taxa.
However, the stratigraphic studies of Frank (1971) and Osborne (1983) suggested that
the sediments may possibly range in age from Miocene to late Pleistocene.
To test this hypothesis, a team from the School of Zoology, University of New South
Wales, commenced stratigraphically controlled collection of bones from several of the
caves. Between 1982 and 1986 bones were collected from Big Sink, the Phosphate Mines,
Bone Cave and Cathedral Cave. The first results of this study have indicated an early to
mid-Pliocene age for the deposits in the Big Sink, hypothesised by Osborne (1983) to be
the oldest stratigraphic units in the complex (Hand et al. 1988). The present paper contin-
ues this study and reports on the sediments and fauna from the floor of the Cathedral Cave.
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 53
DESCRIPTION OF CATHEDRAL CAVE
The Cathedral Cave is the largest of the natural caves in the Wellington Caves
complex. The cave (Fig. 1) consists of a steeply sloping entrance passage which extends
for about 40 metres before expanding into a huge main chamber with dimensions of 45
m long, 20-25 m wide with a domed roof approximately 14 m maximum height. This
chamber now has a nearly level compact earth floor, which is approximately 20 m below
the ground surface. At the northern-most end of the chamber a huge ‘stalagmite’ reaches
from floor to ceiling. The cave derives its name from the resemblance of this flowstone
covered pillar to a cathedral altar.
The chamber has been formed by nothephreatic solution in massive limestone of
the Devonian Garra Formation (Osborne 1983). The walls and roof of the chamber are
largely devoid of speleothems. The chamber has been formed along a fault between thin-
ly bedded limestone which forms much of the roof and eastern walls, and unbedded mas-
sive limestone forming the western roof and walls. The ‘altar’ occurs on this fault line.
Immediately north of the ‘altar’ there is a vertical drop of approximately 3.5 m to the
floor of a second narrower chamber. This floor extends horizontally for approximately 30
m to another vertical drop of 6—7 m terminating in a pond of water. The level of water in
this pond varies with rainfall and the flow in the nearby Bell River. During a flood in
1956 the water rose to a level | m deep throughout the second chamber (1.e. only about
2.5 m below the floor of the main chamber).
The Cathedral Cave (then known as “the Great Cave’) was described and illustrat-
ed by Sir Thomas Mitchell, who first entered this cave and surveyed it in 1830 (Mitchell
1838). Mitchell discovered some fossil bones in Cathedral Cave in 1830, but these were
considered insignificant in comparison to his discoveries in the nearby “Breccia Cave”.
Prior to 1881 extensive bone collections of great significance were made from the
“Breccia Cave” (now known as Mitchell’s Cave) (Dawson 1985). However, there is no
record of any collection from Cathedral Cave during that period.
In 1881 at least two shafts were dug in the earthen floor of the main chamber of
Cathedral Cave during an exploratory expedition undertaken by the Australian Museum
under the direction of the Curator, E. P. Ramsay. The location and depth of these shafts is
indicated by Ramsay (1882), the most productive excavation being Shaft No 2, which
was sunk near the ‘altar’ and described as follows:
‘At No 2 shaft, which we sunk to the depth of 25 feet, many important bones have
been found, and the different floors we went through show that these bones have been
washed in at different periods. In the last layer, the red mud-like breccia which charac-
terised this shaft (at present) has become more sandy, the bones being more perfect here
and less worn’ (Ramsay 1882, p.34).
This tantalising description provided much of the incentive needed to sustain the
most recent excavation of the Cathedral Cave floor, the results of which are described in
this paper.
MATERIALS AND METHODS
The excavation
The position of the UNSW pit, between the “altar” and the south wall of the main
chamber is indicated in Fig. 1. A site close to the location of Ramsay’s “No. 2 shaft” was
preferred because of its promised depth and productivity. Care was taken to avoid the
exact location of the previous shaft. The fact that the cave was continuously in use by
large groups of tourists further limited the choice of sites for the excavation.
Surface dimensions of the pit were 2.0 m x 1.0 m, with the long axis extending
away from the “altar” in a SSE direction. The northernmost edge of the pit was approxi-
mately 1 m from the outer rim of the base of the “altar”, which is delineated by concen-
Proc. LINN. SOC. N.S.W., 117. 1997
54 LATE QUATERNARY SEDIMENTS
tric ridges of flowstone. Surveyed co-ordinates of the pit are Easting 294 294.7 Northing
1 289 237.1 Height 322.6 (Integrated Survey Grid [ISG] of NSW).
Sediments were excavated by hand in ‘spits’, which were of variable depth, rang-
ing from 10 to 20 cm, depending on the nature of the sediment encountered. Each spit
was further divided into 8 horizontal grid segments, labelled A-H. The final depth of the
excavation was approximately 7.25 m from the cave floor surface. All material was
bagged and labelled before being stored in the cave, or removed from the cave for further
treatment, either locally or at the University of New South Wales.
Treatment of the sediments consisted of first drying them, then screen washing
(flyscreen) with water. Residual lumps of sediment which were insoluble in water were
treated further with dilute acetic acid. However, there was generally poor breakdown of
the clay-rich breccia in acid. As far as possible all bone was retrieved and stored for fur-
ther analysis. Other inclusions such as charcoal, gravels, and limestone pieces were also
retained for future examination.
During the excavation samples were collected for detailed sedimentary analysis.
Unfortunately most of these samples were inadvertently destroyed.
Analysis of bone
After screen washing and treatment of the sediments, all bone was collected and
labelled according to the spit and grid sector of origin. Only teeth and jaws have been
used to identify the taxa present in the deposit. The absolute number of individuals per
stratigraphic unit could not be used for comparison between units because each unit rep-
resents an unknown (different) volume of sediment. In order to compare the relative
abundance of various mammalian taxa, the minimum number of individuals (MNI) and
the percentage abundance of each taxon was calculated for each stratigraphic unit. The
estimate of minimum number of individuals was based on a count of identifiable right
and left dentaries and maxillae. The most numerous element was taken as the MNI.
Where a taxon was represented by a few isolated teeth only, it was scored as MNI = 1,
except in the case of the peramelids (bandicoots),which were frequently represented by
isolated molar teeth. These teeth were identified by position in the tooth row (e.g. LM",
RM_)) and the most numerous element was taken as the MNI. The percent abundance of
a taxon in each unit was calculated by the formula MNI /Total MNI per unit X 100. Non-
mammalian taxa were not included in this analysis.
Dental nomenclature follows Luckett (1993). All specimens are registered in the
collections of the Australian Museum, Sydney.
Abbreviations: AM = Australian Museum, QM = Queensland Museum, L = length,
AW = anterior width, PW = posterior width, mm = millimetres.
RESULTS
Stratigraphy
The present floor of Cathedral Cave is nearly horizontal and has been beaten to a
smooth hard surface by tourists over many years. At the site of the excavation the floor
was somewhat irregular and sloped slightly away from the altar. The top metre of soil
showed evidence of extensive disturbance. Fragments of bottle glass, sawn timber, metal,
and even a piece of gramophone record were encountered in Spits 1-3. There was no fur-
ther evidence of disturbance of the sediments by man below Spit 4, approximately one
metre below the present floor. That the sediments beneath Spit 4 were undisturbed was
supported by the presence in Spit 5 of a thin discontinuous layer of flowstone, no more
than 2 cm thick, below which an orange cemented sediment rich in small bone fragments
was encountered.
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE a5)
Over a period of approximately four years the Cathedral Cave excavation was con-
tinued to a final depth of 7.25 m below the present floor of the cave. Throughout the pit
the sedimentary matrix consisted predominantly of heavy red clay. Inclusions varied in
size from large limestone boulders (up to 30 cm in diameter) to gravels, and included both
jagged rock fragments and smooth river pebbles. After consideration of the nature of the
stratigraphy of the deposit, it became apparent that analysis of 10 cm spits was unwarrant-
ed, so samples were pooled for 50 cm intervals for the purpose of faunal analysis.
TABLE |
Stratigraphic description of excavation in floor of Cathedral Cave, Wellington Caves.
Depth (m)
at base of unit
Spit groups
(inclusive)
Description
Phase 3
= 11-5)
Phase 3
—2.0
Phase 3
5)
Phase 3
=A)
4-5
6-10
11-15
16-19
Matrix of heavy red clay mottled with grey loamy textured silt, containing
gravels and limestone nodules of varying size plus occasional small pieces of
flowstone. Residue after washing comprised 30-40% original volume of
insoluble material including bone and calcified clay aggregates.
Bone: low concentration of small fragments of bone, max. fragment length
approx 2cm.
Matrix and inclusions as above. Lens of charcoal in spits 6 and 7 gave cl4
date of 2,950 + 80 BP.
Bone: as above, larger taxa represented by isolated teeth only.
Matrix and inclusions as above. Large boulder approx. 20 cm diameter in spits
11-12. At spits 13-14 there was a hardened ‘floor’ 10 cm thick composed of
indurated clay and flowstone, dipping south at approxmiately 20°. A small
circular area rich in charcoal occurred immediately below this ‘floor’ and gave
a C!4 date of 2540+80 BP.
Bone: As above, max. fragment length approx. 3 cm.
Matrix of heavy red clay, less insoluable residue than overlying strata
including gravels and limestone fragments.
Bone: As above, max. fragment length approx. 4 cm.
Phase 2
=35)
Phase 2
—4.0
Phase 2
ma
20-23
24-28
29-33
Zone comprises a hard ‘floor’ approximately 20 cm thick extending the entire
length of the pit, dipping south at approximately 20° . This consists of heavily
cemented clay with inclusions of limestone boulders and fragments, flowstone
and bone breccia
Bone: Relatively high proportion of bone representing both large and small
taxa. Large bone variably preserved, some weathered prior to deposition,
fragmented. Max. fragment length approx 23 cm. Largest taxa represented by
juvenile individuals.
Matrix of zone immediately below the ‘floor’ comprised unconsolidated red
clay with few large inclusions. At north-west corner of spits 26 A-27A
encountered a limestone boulder >30 cm in diameter with crushed and broken
bone cemented to it. Another very large boulder in 28C. Spit 26 represents top
of a layer of ‘rubble’ approx 30 cm deep dipping to the south as above, but
less heavily indurated than ‘floor’ described above.
Bone: low concentration of small and larger bone fragments which are thinly
encrusted with calcite. Largest taxa represented by juvenile individuals. Max.
bone length = 14cm.
Matrix of heavy red clay, residue after washing comprised from 90% (at top of
zone) to 30% (toward base of the zone) of original volume as insoluble
calcite nodules and limestone pieces.
Bone: Relatively low bone content, small jaws and fragments only, thinly
encrusted with calcite. Max. bone length = 5 cm.
Proc. LINN. SOC. N.S.W., 117. 1997
56 LATE QUATERNARY SEDIMENTS
Phase 2 34-38 Matrix of heavy red clay, residue after washing comprised 30-50% original
volume of small calcite nodules and flakes. Spit 36 A (nth end) contained thin
indurated layer including much small gravel overlying a thin layer of blackened
small bone. Spit 36B encountered top of a limestone boulder approx. 30 cm
diameter.
Bone: Relatively high concentration of large and small bone with largest pieces
approx. 12 cm long; partial macropodid cranium 8 x 11 cm. Bone encrusted
= with red calcite.
Phase 1—5.5 39-42 Matrix of sandy red clay, gravels and occasional large pieces of limestone,
residue after washing containing calcite nodules and flakes. Pooled sample of
charcoal from spit 40 gave a C 14 date of 21,400+700 BP
Bone: Very rich in bone, many larger bones (including robust femur 24 cm
long) , also rich in bones of small taxa. Bone reddish cream in colour, thinly
= 55) encrusted in calcite, not abraded or chewed.
Phase | 43-45 Thin calcite layer on top of spit 43 at north end of pit, petering out towards
south. Matrix of sandy reddish clay with higher gravel and bone content.
Residue after washing contains many flat grey ?calcite flakes.
Bone: Relatively high content of bones of small and larger taxa. Bone well
preserved, cream-brown in colour, no calcite encrustation. Largest bone
fragment 16 cm long. Associated skeletal elements present. Some bone
—6.0 appears gnawed.
Phase 1| 46-50 Matrix of reddish clay with high proportion of sand and gravels. No evidence
of dipping strata. Pooled samples of charcoal from Spit 46 gave C!4 date of
21,350 +1,700 BP. Pooled samples of charcoal from Spit 50 gave C!4 date of
26,800+2000 BP.
Bone: Relatively low bone content, washed clean with water, lacking calcite
encrustation. Bones represent mostly small taxa with larger taxa represented by
isolated teeth only. Largest bone fragment 7 cm long. Some bone slightly
—6.5 gnawed.
Phase 1 51-54 Spit 51 as above. Spits 52 and 53 comprised of grey friable matrix. Thin hard
calcite layer in spit 54, red matrix below this. Charcoal from Spits 51 and 52
gave C!4 dates of 33,800+2000 BP and 32,500+2100 BP, respectively.
Bone: Extremely rich in large and small bone fragments, longest approx. 17 cm;
associated elements present; rich in bones of small taxa. Bone washed clean in
water; colour mottled grey, no calcite encrustation, larger bone not gnawed or
=) abraded.
Phase | 55-57 Matrix red sandy clay, inclusions wash clean in water, not indurated, but
contained many flat grey ‘flakes’ of rock (?calcite ). Many large limestone
boulders encountered below spit 55.
Bone: Relatively high content of creamy coloured bone, washed clean in water,
lacking calcite encrustation; large species represented by isolated teeth and
fragmented bone with highly abraded (? gnawed) ends; largest fragment
= The) approx 9 cm long. No associated elements.
Detailed description of the sediments is provided in Table 1. Stratification within
the sediments was indicated by variations in degree of induration and cementing of lay-
ers to form successive “floors” throughout the deposit. As the excavation proceeded sev-
eral hardened “floors” of varying thickness were encountered as described in Table 1.
Above Spits 30-33 ( -4.25 to -4.5 m) these “floors” dip at a consistent angle of approxi-
mately 20 degrees from the north edge of the pit towards the south, i.e. slope away from
the base of the “altar”, consistent with the hypothesis that they form an extension of the
base of a cone of sediment with its apex under the existing stalagmite called the “altar”.
Below 4.5 m the stratification is nearly horizontal.
The following factors have been considered in formulating a hypothetical interpre-
tation of the history of this deposit:
Proc. LINN. SOc. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 57
e the sequence and direction of slope of hardened ‘floors’ and flowstone layers
e degree of induration of the sediments
e the preservation of bone, including size, colour, fragmentation, association of
elements, presence of juveniles etc.
e the nature of inclusions (other than bone) in the sediments
e the information from C!4 dating of charcoal
Examination of the data presented in Table | has indicated three main phases in the
deposit. These are described and justified below.
Phase |
This is the oldest phase of deposition, and extends from the base of the pit (7.5 m)
to approximately 5.0 m below the surface (Units 55-57 to Unit 39-42, inclusive). The
upper boundary is not delimited by any definite demarcation line; in fact Spit 38 and
Unit 39-42 represents a transition zone. However, Phase | has several characteristics
which define it:
e sediments are approximately horizontal
e predominantly more sandy/gravelly sediments
e much less calcification / induration of sediments than higher levels — almost
completely soluble in water
e bone well preserved, less fragmented, including some associated elements and
intact skeletal elements of larger taxa
e large taxa represented mainly by juveniles
e fauna contains extinct taxa and “megafauna”’
Phase 2
The middle phase of deposition extends from approximately 5.0 m to approximate-
ly 3.0 m below the surface (Units 34—38 to 20-23 inclusive). The upper boundary of this
phase is demarcated by heavily indurated rubble ‘floor’ approximately 20 cm thick.
Characteristics defining Phase 2 are:
e red clay, generally heavily indurated, does not break down readily in water
e large number of boulder-sized inclusions and many smaller rock
e strata dip at approximately 20° from north to south
e large bones highly fragmented, some weathered prior to deposition
e large taxa represented by small jaw fragments or isolated teeth only
e large taxa mainly represented by juveniles.
e fauna contains extinct taxa and “megafauna”
Phase 3
The most recent phase of deposition is represented from -3.0 m to the surface (the
top one metre being disturbed since European settlement). Characteristics of Phase 3 are:
e red clay, variably indurated, mostly breaks down in water
e inclusions consist mainly of gravels, small limestone nodules and small pieces
Proc. LINN. Soc. N.S.W., 117. 1997
58 LATE QUATERNARY SEDIMENTS
of flowstone.
e low bone concentration.
e bone fragments small, large taxa represented by isolated teeth only
e little evidence of stratification — some thin flowstone sheets
e no extinct species represented
TABLE 2
Radiocarbon dates obtained from charcoal excavated from the floor of Cathedral Cave, Wellington Caves.
Asterisk indicates pooled samples. Depths represent depth below present cave floor.
Sample no. Spit Depth Age (yrs BP)
SUA 2097 Spit 6 1.6m 2950 + 80
SUA 2098 Spit 14 2.7m 2590 + 80
ANU 4480 Spit 39 * 5.1m 14,300 + 730
ANU 4479 Spit 40 * 5.2m 21,400 + 700
ANU 4478 Spit 44 * 5.8m 11,900 + 790
ANU 5323 Spit 46 6.1m 21,350 + 1700
ANU 5324 Spit 47 * 6.2 m 28,000 + 1100
ANU 5325 Spit 49 6.4m 23,700 + 1400
ANU 5326 Spit 50 * 6.5m 26,800 + 1100
ANU 5327 Spit 51 6.6 m 33,800 + 2000
ANU 5328 Spit 52 6.7 m 32,500 + 2100
Radiocarbon dating
Charcoal was rare in the Cathedral Cave deposits, but small samples of charcoal
were recovered during the excavation and while screen washing. These were collected in
sealed plastic bags and submitted for radiocarbon dating. The results are presented in
Table 2. The first two samples (Spits 6 and 14) were submitted to the Radiocarbon
Laboratory at the University of Sydney (SUA) in 1984. A further 9 samples (from Spits
39 to 52) were submitted to the Radiocarbon Dating laboratory, Research School of
Pacific Studies, Australian National University (ANU), in 1985 and 1986. In several of
the spits it was necessary to pool small samples of charcoal to obtain a quantity large
enough for dating. Dates derived from pooled samples are indicated with an asterisk in
Table 2.
The irregularities apparent in the sequence of the dates (Table 2) may be due to a
variety of factors, including contamination during handling and pretreatment of small
pooled samples of charcoal, lateral facies changes and variable transportation of different
elements (Archer 1974, Osborne 1984). The date for Spit 44 (ANU 4478) is most likely
to be anomalous and due to younger organic contamination during pretreatment (J. Head,
ANU Radiocarbon Dating Laboratory, pers. comm. 1985).
The C!4 dates presented in Table 2 provide an indication of the probable time span
involved in this deposit, from approximately 34,000 BP (the minimum age of the lowest
sediments) to approximately 2,500 BP for the undisturbed sediments at the top of the
deposit. While there are several inconsistencies in the sequence of the dates throughout
the column, the general trend of the dates indicates that the lower levels of the deposit
are older than the higher levels, i.e. that there is no major stratigraphic reversal present.
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 59
The fauna
Bone fragments were present throughout the deposit (Table 1). The preservation of
the bone varied enormously, ranging from very weathered bone, some crushed bone,
some heavily mineralised well preserved elements and some clean, lightly mineralised
bone. Generally the larger bones were very fragmented and, with few exceptions, large
taxa were represented by small jaw fragments only. No intact jaws or associated skeletal
elements were encountered until more than 5 m below the surface, after which depth the
bone was generally better preserved.
Teeth and jaws referable to marsupial taxa have been identified to species level
where possible. Specific identification of most other non-marsupial remains has not been
attempted, with the exception of some of the rodents, the Ghost Bat, Macroderma gigas,
and the identification of snail shells (Mollusca).
The fauna of the Cathedral Cave deposit is summarised in Table 3, which indicates
the presence/absence data for taxa at each stratigraphic level. At least 38 marsupial taxa
have been identified in the deposit, of which 10 are extinct, and one, Sarcophilus
harrisii, is now confined to Tasmania. Of the remaining 27 extant marsupial taxa only 9
may still occur west of the Great Dividing Range today (Dickman 1994). Of the non-
marsupial taxa, most are rodents, which are abundant in most strata. Microchiropteran
bats are represented in several strata, as are reptiles and small birds.
Table 4 presents the MNI present and % abundance per stratigraphic unit and the
total taxa present. The relative abundance of selected taxa has been graphically depicted
in Fig. 2. The results must be considered in light of the fact that there was considerable
difference between stratigraphic units in the total number of taxa present, thus affecting
comparison of percent abundance values throughout the deposit.
Taphonomy
Preservation of the bone and the observed stratigraphic variations suggest that the
fauna present in the Cathedral Cave floor deposit has complex sources, and it is likely that
several different agents were responsible for the accumulation of bones. In each strati-
graphic unit of this deposit at least 85% of the animals present represent taxa with average
body weights of less than 1.5 kg. Bones of small animals (less than 1.5 kg in body weight)
are most likely to be derived from owl pellet deposits or the prey of Ghost Bats (Baird
1991), while the remains of larger animals may be accumulated by carnivores or scav-
engers or accumulate because the cave acts as a natural pit-trap. Analysis of the relatively
intact jaw fragments of larger animals in the deposit has revealed that 88% of the macrop-
odines present are juveniles, only three specimens representing adult individuals.
Carnivores and scavengers are extremely rare in the deposit (4 jaws only), but in each case
these are from adult individuals. These data indicate that large animals mainly entered the
cave by falling in, their remains attracting scavengers, some of which also succumbed.
It is considered that the three phases recognised in this study represent periods in
which there were different agents of accumulation or sources of the bone.
Phase 1 has the characteristics of an attritional entrance facies deposit (Baird 1991).
This profile is characteristic of the distal elements of a talus cone, not markedly reworked
since deposition (associated skeletal elements). Large taxa are represented by juveniles only,
except for scavengers and carnivores. This is characteristic of a pitfall trap. The abundance
of small mammals is consistent with accumulation due to owls and ghost bats. Within Phase
1 stratigraphic horizons are indistinct, with the exception of the unit represented by Spits
51-54 which was grey in colour, rather than the otherwise predominantly red sediments.
However, the composition of the fauna varied considerably from spit to spit. For example,
Spits 51-54 contained the greatest number of taxa, but fewer individuals than units above
and below it (Table 4). Spits 51-54 also contained the highest fraction of bone representing
large species (Fig. 3), despite the fact that it also contained the largest number of Ghost Bats.
Proc. LINN. SOc. N.S.W., 117. 1997
60 LATE QUATERNARY SEDIMENTS
TABLE 3
Faunal list from the floor of the Cathedral Cave, Wellington Caves, indicating presence/absence of each taxon
per strtigraphic unit of the excavation
Spits 4-5
Spits 6-10
Spits 11-15
Spits 16-19
Spits 20-23
Spits 24-28
Spits 29-33
Spits 34-38
Spits 39-42
Spits 43-48
Spits46-S0
Spits 51-54
Spits 55-57
Sminthopsis crassicaudata —
Sminthopsis murina —
Sminthopsis sp indet eT
Antechinus sp cf A. flavipes — | Sp re alin ear la ae ee
Phascogale tapoatafa — ee
Phascogale calura ne eae
Dasyurus viverrinus — fs
Dasyurus geoffroii — — | — Se eee alee a
Dasyurus hallucatus — |—
Sarcophilus harrisii eae
?Dasyuroides sp. = tes
Perameles gunnii Ss eae ee ape) ae) ai ata tS See A 2
Perameles nasuta Se SS a eS es eae ee eS eS
Isoodon obesulus — | — EAE ee PE Pipe ec Ne | RR | ei (pee
Thylacinus cynocephalus — a—
Trichosurus sp. —|/|— ee el
Petaurus sp. cf P. breviceps a (ee
Pseudocheirus sp. sites
Acrobates pygmaeus —|—
Aepyprymnus rufescens =— — | —}] — — }— |] —
Bettongia sp —
Potorous tridactylus — | — —
Macropus giganteus — —
M. (Osphranter) sp. —
M. (O) altus # a
M. sp.cf. M. (N.) agilis # #
M. sp. cf M. (N.) dorsalis — —
M. sp. cf. M. (N.) rankeni #a #
M. (M.) titan # # # #
M. (M.) ferragus
Protemnodon sp. — _
Petrogale sp. # # — # #
Thylogale sp. "| # Ed a
Onychogale sp. | ieee
Simosthenurus oreas | — # #a #
Diprotodon sp. | x Xx
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 61
S fy Sef SS SB ea Sol Se ee ta
— Kr d d N
Ta = ] ] | | | | af yl “fp |
ali | = Ne) i) t ony t D sa) Ve) = wn
\o — — N N N (oe) ioe) + + ra) ra)
Nn Nn n n Nn n Nn Nn N Nn Nn N N
a=] = = = = = = = = = = | = =
Q a Qu a a ioe ioe a QO. i= Oo. jor Qu
N N N nN Nn Nn Nn nN nN Nn YN N N
Thylacoleo carnifex
Vombatus sp.
Pal
*
Pal
Phascolarctos sp ae pepe
Rodent indet (med) | SS RB es MIT |e lh a a cate eer eg ete i
Rodent indet (sm) — — — = oe pes sotal at pas pul hie
Mastacomys fuscus | ie a pei
Conilurus albipes SPS felaetleahael alae pal Sha) aH
Hydromys sp. ra
Small bats indet —|— — ar ee
Macroderma gigas eee ee foe
Varanus sp. — =e
Teliqua sp. —|— ast
Small reptiles — _ — fee aes
Small birds indet —= =) || = etn yy aes
Mollusca (snails) — pas ie ee ee =
Legend: a = associated elements; # = juvenile; x = enamel fragment only; bold italics = extinct.
In Phase 2 a high degree of disturbance is suggested. The large number of lime-
stone boulders and rocks, the highly fragmented nature of bone, and the complete
absence of any associated skeletal elements, supports an hypothesis that the bones and
sediments have been redeposited here. The sediments of this phase vary in the degree of
cementation and include two distinct hardened “floors”, each approximately 20 cm deep.
The consistent dip of the strata toward the south away from the present ‘altar’ suggests
that the source of the sediments is likely to be from a point above the ‘altar’. The total
number of taxa and the number of individuals represented is much lower than in Phase 1
(Table 4). The proportion of larger species is also higher than in Phase 1 (Fig. 3). The
predominant presence of juveniles among the large fauna indicates that the original
deposit acted as a pitfall trap for large animals. Ghost Bats are absent from the fauna of
this Phase and it is likely that small mammal remains are derived from owl pellets.
There is little evidence in the upper phase (Phase 3) to suggest the probable source
of inclusions. There are no large bone fragments present, and large species are represent-
ed by isolated teeth only. This differs markedly from the earlier phases and indicates that
the entrance to the cave was restricted. The entrance site above the ‘altar’, suggested to
have been present during Phase 2, had apparently closed by this time, as there is no evi-
dence of large rubble or characteristics of pit-fall accumulation.
It is unlikely that this deposit was an active vadose environment at any time during
the period of deposition (i.e. during the last 35,000 years), so water transport 1s not likely to
be responsible for the accumulation of the bones. The highest terraces of the Bell River are
considerably lower than the present cave entrance, and probably date from the Pliocene
(Francis 1973) and the cave has never been a vadose cave, although the action of ground-
water may have contributed to the secondary distribution of the elements in the deposit.
However, it is likely that level of groundwater has intermittently risen to at least
the level of the present floor of Cathedral Cave over the period represented by the floor
deposits (note 1954 flood level in lower chamber). This intermittent saturation and dry-
Proc. LINN. SOc. N.S.W., 117. 1997
62 LATE QUATERNARY SEDIMENTS
TABLE 4
Faunal list from Cathedral Cave, Wellington Caves, indicating the Minimum Number of Individuals (MNJ) and
percentage abundance of each taxon per stratographic unit
Spits 6-10
Spits 11-15
Spits 16-19
Spits 20-23
Spits 24-28
Sminthopsis crassicaudata
Sminthopsis murina
Sminthopsis sp. indet 1
Antechinus sp. cf. A. flavipes
Phascogale tapoatafa
Phascogale calura
Dasyurus viverrinus
Dasyurus geoffroii
Dasyurus hallucatus
1
1
Sarcophilus harrisii
?Dasyuroides sp.
Perameles gunnii
Perameles nasuta
Isoodon obesulus
Thylacinus cynocephalus
Trichosurus sp.
Petaurus sp. cf P. breviceps
Pseudocheirus sp.
Acrobates pygmaeus
RePwWh
Aepyprymnus rufescens
Bettongia sp
Potorous tridactylus
Macropus giganteus
M. (Osphranter) sp.
M. sp. cf. M. (O.) altus
M. sp.cf. M. (N.) agilis
M. sp. cf. M. (N.) dorsalis
M. sp. cf. M. (N.) rankeni
M. (M.) titan
M. (M.) ferragus
Protemnodon sp.
Protemnodon brehus
Petrogale sp.
Thylogale sp.
Onychogale sp.
Sthenurus sp. cf. S. oreas
Diprotodon sp.
Thylacoleo carnifex
Vombatus sp.
Phascolarctos sp.
Rodent indet (med)
Rodent indet (sm)
Mastacomys fuscus
Conilurus albipes
Hydromys sp.
Small bats indet
Macroderma gigas
Varanus sp.
Teliqua sp.
Small reptiles
Small birds indet
1
1
1
Total MNI Non-volant mammals| 27 63
TOTAL TAXA INCLUDED | 12 17
Proc. LINN. Soc. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 63
foe) co NN wy eS wt ™
of ae at Bt f wie
OD wr oO (oa) \o iomal a)
N lon) foe) wt wt Ww wy
8
7.9 8 4
3
B2 0 4
2 1
5 2
1
1.6 3
1
3.2 4
4.8 1
1.6 1
1
1.6 2
1
1.6
1.6
1.6 1
1.6
1
1.6 1
1
1
1
1
1.6 1
1
55.6
6.3
1.6
3)
Proc. LINN. SOc. N.S.W., 117. 1997
0.6
0.3
0.3
64 LATE QUATERNARY SEDIMENTS
ing of the sediments would account for the succession of calcified ‘floors’ and flowstone
layers encountered in the deposit. If the water level were to rise above the level of the
sediments (as is likely) this would account for the levelling of the cave floor.
A high level of induration and calcification, as is observed in Phase 2 sediments, could
result from a long period of stagnation of calcite-rich water — or a prolonged dry period fol-
lowing after a period of saturation.
SYSTEMATIC ANALYSIS
Mammalia
Marsupialia
Dasyuridae
Larger dasyurids (body weight greater than approximately 1—1.5 kg) are represent-
ed by three species of Dasyurus and the devil, Sarcophilus harrisii, all of which are scay-
enger / carnivores.
Species of Dasyurus have been identified from jaw fragments by comparison with
specimens from the modern fauna in the Australian Museum. In the case of maxillary
fragments, identification was assisted by the comparison of dental measurements. The
ratio of protocone-parastyle length / protocone-metastyle length of M4 distinguishes
between D. geoffroii and D. hallucatus. D. viverrinus is distinguished by being larger
than the other two species.
In Phase | the Western Quoll, D. geoffroii is represented in all units by well pre-
served jaws. The Eastern Quoll, D. viverrinus, is also represented in units 51-54 by well
preserved jaws. The Devil, Sarcophilus harrisii is represented by a single intact right
mandibular ramus. Dental measurements of this specimen place it in the range of overlap
in size between the largest individuals of the modern Tasmanian population and the
smallest individuals attributed to the Pleistocene species, S. laniarius (Dawson 1982a).
Given that this specimen comes from the oldest strata of the Cathedral Cave deposit,
associated with C!4 dates of approximately 33,000 BP, it is possible that it represents the
Pleistocene species, however, this diagnosis cannot be made without a larger sample.
In Phase 2 Dasyurus geoffroii 1s represented in all but one of the units in this
phase. In Spits 34-38 both D. geoffroii and D. hallucatus (the Northern Quoll) are pre-
sent, while a single maxillary fragment appears to represent D. hallucatus in Spits 29-33.
In Phase 3 large dasyurids are extremely rare, Dasyurus viverrinus and D. geoffroii
being represented by a few isolated teeth only.
The most common species of Quoll represented in the Cathedral Cave deposit is D.
geoffroii. This species was present in the western division of New South Wales at the
time of European settlement, but has not been captured in New South Wales since 1857
(Dickman 1994). D. viverrinus has not been recorded in western New South Wales in
historic times, being confined to coastal regions of south-eastern Australia and Tasmania
(where it is still extant).
At least five species of dasyurid smaller than 1.5 kg body weight have been identi-
fied, all of which are extant in the modern fauna of eastern Australia. At least two
species of Sminthopsis are represented, S. murina and S. crassicaudata, with the former
being almost ubiquitous, while S. crassicaudata is rare, being present in two widely sep-
arated levels (Phase | and Phase 3) only. Antechinus sp. cf A. flavipes is present below
Spit 20. Phascogale tapoatafa is present in Phase 1 and Phase 2, while P. calura is pre-
sent in the lower spits of Phase | only.
The greater relative abundance of small dasyurids in Phase 1 compared with higher
levels of the deposit is marked. Both A. flavipes and S. murina show dramatic peaks in
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 65
90
20> Sminthopsis murina Large rodents
Antechinus flavipes 60
ie Phascogale tapoatafa
30
10 Perameles nasuta
0
® ® 6
© 107 Perameles gunnii E
= g & 697 Small rodents
(‘Ss Cc
= =)
a co}
(9) oO
32 32
** 10> jsoodon obesulus r
40
10> Dasyurus geoffroii
20
10> Aepyprymnus rufescens
10-4 Mastacomys fuscus
10> Acrobates pygmaeus
0
° 207 Conilurus albipes
104 Petaurus breviceps
TpHases! pHase2! puHaser | TpHases ! pHase2! pHase1 |
3.0 5.0 7.5 3.0 5.0 7.5
Depth (m) Depth (m)
Figure 2. Relative abundance of selected mammal taxa represented in the sediments from the floor of Cathedral
Cave, Wellington Caves. Data from Table 4.
Proc. LINN. Soc. N.S.W., 117. 1997
66 LATE QUATERNARY SEDIMENTS
Percent abundance of small mammals
Ww
I
wT
%
ee
a
w
Spits 6-10
Spits 11-15
Spits 16-19
Spits 20-23
Spits 24-28
Spits 29-33
Spits 34-38
Spits 39-42
Spits 43-45
Spits 46-50
Spits 51-54
Spits 55-57
Figure 3. Relative abundance of small mammal taxa with body weight less than 1.5kg in the sediments from
the floor of Cathedral Cave, Wellington Caves. Data from Table 4.
the lower units which contain the Ghost Bat, Macroderma gigas. These spits also contain
the greatest diversity of dasyurid taxa in the deposit. In fact small dasyurids represent
nearly 35% of the total number of individual animals recorded from Spits 51-54 (Table
4). While the foraging habits of the Ghost Bat most likely account for the abundance of
individuals in that stratigraphic unit, other factors may account for the rich species diver-
sity in the lower spits of Phase 1. The high diversity of small dasyurids in the
‘Macroderma’ spits of Phase | coincides with an increase in relative abundance of small
rodents and a sudden sharp decrease in relative abundance of larger rodents (Table 4, Fig.
2). Detailed identification of the rodent taxa present must be undertaken before hypothe-
ses to explain these observations can be developed.
All the species of small dasyurids identified in the Cathedral Cave deposit (with
the possible exception of Phascogale calura and ?Dasyuroides sp.) have ranges in the
modern fauna which encompass the Wellington Valley, at least peripherally (Strahan
1983). Similarly, all species are common components of other late Pleistocene fossil fau-
nas of eastern Australia (Lundelius 1983). With the exception of Sminthopsis crassicau-
data and Phascogale calura, which have predominantly xeric habitat preferences, the
Species present are characteristic of woodland habitats, and none has particularly rigid
vegetation or climatic limitations, although microhabitat preferences vary between
species (Strahan 1983). Fox (1982) suggests, rather, that species separation is effected by
body size and life strategies in small dasyurids, rather than climatic limitations.
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 67
Thylacinidae
Thylacinus cynocephalus is represented in Phase 2, Spits 34-38, by an isolated
upper molar only. The only other occurrence is the presence of two left mandibular rami
(adult) referable to TZ. cynocephalus in Phase 1, Spits 51-54. The dental dimensions of
these specimens fall in the upper size range of modern T: cynocephalus from Tasmania,
and within the range of Pleistocene populations from Wellington Caves and Naracoorte
Caves (Dawson 1982b).
Peramelidae
Throughout the deposit peramelids are represented by jaw fragments which rarely
retain the molar teeth, but isolated teeth are very common. Isolated teeth have been identi-
fied as to position in the tooth row and the estimate of minimum numbers of individuals has
been based on the counts of teeth according to the method described earlier in this paper.
Isoodon obesulus is present throughout Phase | and Phase 2 of the deposit, increas-
ing in relative abundance in Phase 2. In Phase 3 I. obesulus occurs in the uppermost units
only. Two species of Perameles are also represented throughout the deposit. P- nasuta and
P. gunnii were identified by comparison with jaws and dental characteristics of skulls
from the modern fauna in the collections of the Australian Museum, Sydney, and with the
descriptions and measurements given by Freedman (1967). P. nasuta is the most abundant
peramelid taxon throughout. Both species of Perameles are less abundant in Phase | than
in higher levels, and the relative abundance of P. nasuta over P. gunnii is slightly greater
in Phase 1. In Phase 2 the relative abundance of both species of Perameles shows near
parallel fluctuations. In Spits 16-19, at the base of Phase 3 both P. nasuta and P. gunnii
show a sharp increase in relative abundance, at the same time that /. obesulus temporarily
disappears from the fauna. Subsequently, in the upper units of Phase 3, P. gunnii is more
abundant and more consistently present than P. nasuta. (Fig. 2).
The lower relative abundance in Phase | of all bandicoots, and especially of
species of Perameles, coincides with sharp increases in the relative abundance of small
dasyurid taxa and small rodents (Fig. 2). These trends probably reflect the food prefer-
ences of the Ghost Bat, Macroderma gigas, which is the most likely agent of accumula-
tion of small mammals in the earliest phase of deposition.
Although these three bandicoot species have not previously been identified in the
same deposit, their occurrence together is not incongruous. Gordon and Hulbert (1989)
cite these species (plus J. macroura) as characteristic of the peramelid fauna of the
coastal/sub-coastal southern humid zone, extending in part into the semi-arid zone of
eastern Australia. This contrasts with the species characteristic of true semi-arid areas
(including J. auratus, P. bougainville, P. eremiana, Chaeropus sp. and Macrotis sp.),
none of which have been identified from the Cathedral Cave fauna.
Gordon and Hulbert (1989) note that the habitat requirements of bandicoots are
extremely flexible and opportunistic, although each species may have a preferred habitat.
They suggest that P. nasuta and P. gunnii may exhibit competitive exclusion in their
modern distribution (e.g. in Tasmania P. gunnii thrives in the absence of P. nasuta).
However, J. obesulus occurs with P. gunnii in Tasmania and with P. nasuta throughout
their ranges in the modern mainland fauna and in most late Pleistocene faunas from east-
ern Australia which contain bandicoots. All three species appear to prefer open
forest—grassland habitat.
This is the northernmost record of the occurrence of P. gunnii and this species was
not recorded from the Wellington area at the time of European occupation. P. gunnii is
known from mainland late Pleistocene faunas of South Australia ( Smith 1972, Pledge
1990) and Lake Victoria in western New South Wales (Marshall 1973) as well as from
the modern fauna of Victoria and Tasmania (Gordon and Hulbert 1989).
Proc. LINN. Soc. N.S.W., 117. 1997
68 LATE QUATERNARY SEDIMENTS
Phascolarctidae
Phascolarctos cinereus is represented in Phase 1 by a small mandibular fragment
(Spits 51-54) and several isolated molars (Spits 46 and 51—54). In the modern fauna this
species inhabits wooded areas and river channels throughout New South Wales, its distri-
bution being dependent on the presence of suitable food trees rather than climatic factors.
Thylacoleonidae
Thylacoleo carnifex is represented in Phase 2 by an isolated left I, in Spits 24-28
and another isolated left I, in Spits 34-38. The species is represented in Phase | by a left
mandibular ramus in Spits 39-42, and by an isolated P in Spits 55-57. Well preserved
remains of Thylacoleo are common in museum collections from Cathedral Cave (Dawson
1985). Many of these were collected by Henry Barnes in 1881 (Ramsay 1882), but unfortu-
nately the original records do not include precise stratigraphic locations for the specimens.
Acrobatidae
Acrobates pygmaeus occurs in Phase | only, and then only in Spits 46—54, coinci-
dent with the Ghost Bat, which is most likely to be the accumulating agent. In the mod-
ern fauna, A. pygmaeus has a wide range of climatic tolerance, and inhabits eucalypt for-
est throughout the eastern coast, ranges and slopes. It is therefore unlikely that its limited
distribution in the Cathedral Cave deposit is the result of climatic or vegetation factors.
Phalangeridae
Trichosurus vulpecula is represented by isolated molars in the upper spits of Phase
3. It has not been collected from the Phase 2 levels of the Cathedral Cave floor deposit,
but is represented in all spits of Phase | below Spit 46 and by associated left and right
mandibular rami from Spits 51-54. T. vulpecula is a generalist in its dietary preferences
and inhabits a wide range of habitats. The species is common in the Wellington area
today, and its absence from the Phase 2 levels of the Cathedral Cave is probably due to
unknown taphonomic factors rather than indicating its absence from the fauna of the area
during this phase of deposition.
Petauridae
Petaurus breviceps occurs only in the upper spits of Phase 1 (in relatively low
abundance) and at the top of Phase 2 / base of Phase 3, where there is a sharp peak in
abundance of this species (Table 4, Fig. 2). Occurring as they do at times of transition in
the deposit, it is tempting to hypothesise that some factor related to those transitions
could account for the disjunct distribution of this species in the deposit. It is most likely
that they represent the prey of owls, since the species is not represented in the
‘Macroderma’ spits at the base of Phase 1. Given the modern range of P. breviceps,
where it inhabits forested areas throughout the entire east coast of the continent, extend-
ing inland to the western slopes, it is unlikely that climatic limitations are responsible for
the observed distribution. P. breviceps has been recorded from late Pleistocene faunas
through the eastern margin of the continent, extending from Russenden Cave,
Queensland (Archer 1978) to the Naracoorte region of South Australia (Pledge 1990)
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 69
Pseudocheiridae
Pseudocheirus sp. is extremely rare in the deposit, being represented by a single
maxillary fragment at the base of Phase 2. Ringtail possums are not uncommon in other
late Pleistocene faunas, and occur in the Wellington region in the modern fauna, so their
virtual absence in the Cathedral Cave deposit is difficult to explain.
Potoroidae
Potoroinae
Three potoroine species are represented sporadically through the deposit. The most
common, Aepyprymnus rufescens, occurs at low abundance at the base of Phase 1, peaks
suddenly in the upper levels of Phase 2 and Phase 3, but is absent from the intervening
levels (Table 4, Fig. 2). Potorous tridactylus 1s represented by a single mandibular frag-
ment in each of Spits 51-54, Spits 39-42 and in Spit 38. Its distribution is thus effective-
ly restricted to Phase 1 of the deposit. Bettongia sp. is represented by two isolated pre-
molars, also in Phase 1. Aepyprymnus rufescens inhabits well grassed open forest, and is
now almost entirely confined in its distribution to coastal and southeastern Queensland,
with a relict population on the Murray River in NSW (Strahan 1983:190). A record of
this species from the late Pleistocene fauna of Henscke’s Cave in South Australia (Pledge
1990) suggests that its original range extended throughout mesic areas of eastern
Australia, although it occurs in few fossil faunas of the late Pleistocene.
TABLE 5
Dental dimensions (mm) of Macropus titan from Cathedral Cave, Wellington Caves. a = approximate.
AM F69889_ AM F69890 AM F69891 AM F69892 AM F69893 AM F69894
M2 L 14.5a
PW 12.6
M3 IL 15.8
PW 12.8
P> L 8.8 8.3 8.19.0
PW 6.0a 4.6 4.8 5.1
dP3 IL 11.8 10.8 11.0 10.8 11.0
PW Ved) esa 6.9 7.0 Fol
Occlusal length B = 16.0 mm (F69888)
Proc. LINN. Soc. N.S.W., 117. 1997
70 LATE QUATERNARY SEDIMENTS
Macropodidae
Macropodinae
Macropodine species are represented in all units of Phase 1 and Phase 2 of this
deposit. However, in Phase 3 only a few isolated teeth have been recovered, most refer-
able to Macropus giganteus. This undoubtedly reflects the taphonomy of the Phase 3
deposits, since it would be expected that the larger kangaroos of the modern fauna were
present in the Wellington region throughout the Holocene period.
Five specimens from Phase 2 and two from Phase | are referable to Macropus
titan. All jaw fragments represent juvenile individuals. Dimensions of these specimens
are given in Table 5. Comparison with published data for populations of M. titan from
Queensland and Lancefield (Bartholomai 1975, Dawson 1982c), and with data for the
old collections from Wellington Caves (Dawson 1982c) indicates that the specimens all
fall well within the size range of the Pleistocene species, M. titan, rather than the smaller
M. giganteus .
A single mandibular fragment (juvenile) from Phase 2 (Spits 24—28) represents the
giant kangaroo, Macropus ferragus. Dimensions of the partially erupted My of F69895
are: length = 19.5mm, anterior width = 11.8mm. This species is present in the old collec-
tions from Wellington Caves and the recent collections from Bone Cave, and otherwise
recorded from Pleistocene deposits of the eastern Darling Downs, Queensland, and Lake
Victoria and Lake Menindee in western New South Wales (Dawson and Flannery 1985).
The Pleistocene Wallaroo, Macropus (Osphranter) altus, is represented by two
specimens, a juvenile maxillary fragment from Phase 2 (Spits 24—28) and left and right
maxillae of one adult individual from Phase 3 (Spits 46-50). Although morphologically
identical to M. robustus, the modern wallaroo or euro, M. (O.) altus is considerably larg-
er. Dental dimensions of the Cathedral Cave specimens agree in size with the holotype of
M. (O.) altus from the eastern Darling Downs, Queensland and with a larger sample from
the old collections from Wellington Caves (Dawson 1982c). A species of M. Osphranter
is also represented by three isolated teeth, an I), P3 and an upper molar, in Spits 51-54,
and two isolated upper premolars in Spits 55—57.
Several species of the Macropus wallabies (M. Notamacropus spp.) (Dawson and
Flannery 1985), are present in the deposit. In Phase 2 M. cf. M. (N.) dorsalis is represent-
ed by two juvenile mandibles in Spits 20—23, while a jaw of a juvenile M.cf M. (N.)
agilis occurs in Spits 24—28. In Phase 1 two juvenile mandibles represent M. cf. M.(N.)
dorsalis (Spits 43-45) and a single jaw fragment is referable to M. cf M. (N.) agilis (Spits
51—54). Neither of these taxa have occurred in the Wellington region in historic times,
being now confined to northern Australia. However, the larger Pleistocene form of M.
(N.) agilis, M. (N.) a. siva, is relatively common in old collections from Wellington
Caves (Dawson 1985) and has been reported from Pleistocene deposits in Victoria
(Marshall 1974) and New South Wales (Marshall 1973, Gorter 1977). Macropus dorsalis
has been reported from the Pleistocene Lancefield Swamp fauna of western Victoria
(Gillespie et al. 1978) but was not identified in the old collections from Wellington
Caves (Dawson 1985).
A new species of large wallaby referable to Macropus (Notamacropus) is present
in Phase | of the deposit. Remains similar to (nis species have been recorded, but not yet
formally described, from the old collections from Wellington Caves (Dawson 1982c and
1985, as Macropus rankeni n.sp.) and from the recent collections from Bone Cave
(Dawson in prep). Associated elements from Spit 43 represent the upper and lower denti-
tion, and limb and pedal bones of a juvenile individual. The specimens from Cathedral
Cave vary from the Bone Cave material in premolar morphology and in being slightly
smaller, and may represent a ‘dwarfed’ version of the Bone Cave species. This species
may have its closest affinities with Macropus dorsalis, but varies considerably from that
species (and from all other Notamacropus wallabies), in being much larger, having a rel-
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 71
atively longer, more slender diastema and in premolar morphology. The elongate
diastema and relatively high crowns of the molars indicate that this was a grazing animal.
Large extinct wallabies of the genus Protemnodon are extremely rare in the
deposit, being represented by isolated teeth in Spits 29-33 and Spits 55-57 only. The
teeth represented in the present collection are closest to P. brehus in size, but there is
insufficient material for definite specific determination. Species of Protemnodon are well
represented in the old collections from Wellington Caves but it 1s unlikely any of those
specimens came from the Cathedral Cave (Dawson 1985).
Rock wallabies (Petrogale sp) are represented in Phase | and Phase 2 by jaw frag-
ments of juveniles. Although Petrogale sp. does not occur in Phase 3 of the deposit, the
Wellington region falls within the modern range of P. penicillata, which inhabits suitable
rocky areas in sclerophyll forest of inland New South Wales (Strahan 1983:211).
The presence of species of Thylogale and Onychogalea in the lower levels of the
Cathedral Cave sediments_has been determined on the basis of isolated teeth (the distinc-
tive posteriorly grooved I~ of Thylogale sp and the small narrow I, of Onychogalea sp.)
The presence of these taxa together in the lower levels of Phase | has contradictory eco-
logical implications. O. fraenata inhabited shrubland and grassy woodland of the western
slopes and plains last century (Strahan 1983:205), but all extant species of Thylogale are
today confined to densely forested habitats of the east coast and ranges.
Sthenurinae
This sub-family of extinct macropodids is represented by 6 specimens from Spits
below 6 metres. An isolated I’ is the only occurrence in Phase 2 of the deposit (Spits
34—38). In Phase | a juvenile left maxillary fragment was found in Spits 43-45, associat-
ed left and right mandibular rami of a juvenile individual in Spits 51—54, and three isolat-
ed teeth (P3, I; and M>) in Spits 55-57. Dental dimensions of these specimens are given
in Table 6. Identification of these fragments as Simosthenurus oreas is based on morpho-
logical comparison of the mandibular teeth (AM F69880, F69881, F69883 and F69884)
with a cast (AM L1728) of the holotype, QM F2923, from the Darling Downs in
Queensland, and with the descriptions of Tedford (1966). S. oreas is a poorly known
species and the upper dentition has not previously been described. The maxillary frag-
ment, AM 69885, has the following dental dimensions ps L = 15.3mm, AW = 8.0mm,
PW = 9.8mm; dP” L = 9.5mm, AW = 8.6mm, PW = 9.8mm; MIL = 11.2mm, AW =
11.0mm, PW = 11.2mm. The teeth of this fragment are in the same size range as the
lower molars of S. oreas, although size alone would not necessarily distinguish this
species from similarly sized Sthenurus andersoni or Simosthenurus orientalis. The max-
illary fragment AM F69885 has therefore been tentatively assigned to S. oreas pending
comparison with more confidently assigned maxillary dentition of this species.
Three species of Simosthenurus and 2 species of Sthenurus have previously been
recorded in the Old Collections from the Wellington Caves (Dawson 1985). As far as can
be determined from their documentation, none of the specimens from the old collections
came from Cathedral Cave. It is most likely that the three specimens of S. oreas in the
old collections came from the Phosphate Mine, since two of the three specimens were
originally part of the collections of the NSW Mining Museum, and the third bears docu-
mentation indicating it was collected in the “drives put in ..... by a Phosphate Company
in search of fertiliser” (Anderson 1932). S. oreas is present in the new collections from
Bone Cave, but is otherwise only recorded from the Pleistocene deposits of the eastern
Darling Downs, and from Bingara in northern New South Wales (Tedford 1966). All
species of Sthenurus and Simosthenurus represent browsing animals. Tedford (1966) has
suggested that Simosthenurus, with its deep skull, short snout and heavily ankylosed
jaws possibly browsed woodland vegetation rather than shrubs.
Proc. LINN. Soc. N.S.W., 117. 1997
7 LATE QUATERNARY SEDIMENTS
TABLE 6
Dimensions (mm) of the cheek teeth of specimens referable to Simosthenurus oreas from Cathedral Cave,
Wellington Caves.
AM F69880 AM F69882 AM F69883 AM F69884 QM F2923*
P5 E 10.0
AW 6.8
PW 8.0
P3 Ie 14.0 14.3
AW 6.0 6.4
PW 8.5 =
dP3 i 10.2
AW 9.0
PW 9.0
M IL 13.2 11.8
AW 10.5 9.8
PW 10.7 10.0
My L 14.9 14.8
AW 11.5 11.5
PW 11.9 11.6
I Depth 12.0% 18.2
* measurements taken from a cast (AM L1728) of the holotype.
** tooth not fully erupted.
Other marsupial taxa
Both Vombatidae and Diprotodontidae are represented in the deposit. However no
analysis of these groups has been attempted, since they are represented only by small
tooth fragments or fragments of dental enamel which are too incomplete to allow identi-
fication to genus level.
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE Ws
Placentalia
Rodentia
Rodents are by far the most abundant animals represented in the Cathedral Cave
deposit, comprising in total more than 70% of the individuals present in all except three
spits (Table 4). This study has not included detailed analysis of the rodent fauna, and
analysis at species level has been confined to two taxa only, Conilurus albipes and
Mastacomys fuscus.
C. albipes occurs throughout the deposit, at relatively low levels of abundance in
Phase 1, increasing to moderately high levels in the top of Phase 2 and middle of Phase 3
(Table 4, Fig. 2). Now extinct, C. albipes occurred in the Wellington area early last cen-
tury, and has been recorded from late Pleistocene faunas of mesic eastern Australia from
Queensland (Russenden Cave, Archer 1978) to South Australia (Henscke’s Cave, Pledge
1990). According to historical records it had a semi-arboreal habitat, and nested in low
hollow tree branches (Strahan 1983:382).
Mastacomys fuscus has a disjunct distribution in the Cathedral Cave deposit, being
present only in the base of Phase 1, and the middle spits of Phase 2 and Phase 3, respec-
tively (Table 4, Fig. 2). M. fuscus is abundant in the late Pleistocene fossil faunas of the
southeastern highlands such as the faunas from Jenolan Caves (Morris et al., this vol-
ume) and Wombeyan Caves (Hope 1982). It is also present in the late Pleistocene of the
Naracoorte region (Victoria Cave and Henschkes Cave, Pledge 1990) and in the Seton
Rock Shelter on Kangaroo Island (Hope et al. 1977). The distribution of M. fuscus in the
modern fauna suggests a preference for cool moist climatic conditions and dense ground
cover (Seebeck 1981). The occurrence of this species in the middle of Phase 2 corrobo-
rates other evidence of a cooler climate in the Wellington region at that time, but the
occurrence in Phase | appears to be anomalous on climatic grounds because of the asso-
ciated presence of the Ghost Bat (which implies a warmer climate).
Chiroptera
Microchiroptera
Unidentified microchiropteran bats are represented in the lower spits of Phase 1,
where they probably formed part of the diet of the Ghost Bat. They appear to be absent
from most of Phase 2, with the exception of the upper spits in the transition zone to
Phase 3, where they again occur in the middle spits of this zone. This distribution possi-
bly reflects changing entrances to the cave.
Megadermatididae
The Ghost Bat, Macroderma gigas, is represented in the lower levels of Phase 1,
its relative abundance peaking sharply in Spits 51-54, where it comprises 45% of the
individuals present (Table 4, Fig. 2). Coincident with this peak, the sediments changed
from predominantly reddish to grey, and positive tests for the presence of phosphates in
the sediments indicated a high proportion of guano (G. Hodge, pers comm, 1991). The
data suggest that at this period the cave was most probably the home of a breeding
colony of Ghost Bats.
Non-mammals.
Small reptiles and small birds are represented sparsely throughout the deposit (Table
4). A species of Varanus is represented at the base of Phase | and the top of Phase 2 only.
Of non-vertebrate remains, 11 shells of terrestrial snails (Mollusca) were found in
the sediments and identified by Dr J. Stanisic of the Queensland Museum, as follows: In
Phase | Spit 53, Nevistitis aridorum, Spit 43, Charopid sp.; Phase 2, Spits 30-40, N. ari-
Proc. LINN. Soc. N.S.W., 117. 1997
74 LATE QUATERNARY SEDIMENTS
dorum, Galidistes sp. Elsothera sp. ; Phase 3, Spit 14, Galidistes sp, Glyptopupoides
egregia. With the exception of the charopid species from Phase | all these species are
found in the area today and are quite characteristic of limestone outcrops of the central
west of New South Wales. The charopid, however is typically found in wetter forests of
eastern New South Wales (J. Stanisic, pers comm. 1991).
DISCUSSION
Osborne (1984) has emphasised the great difficulties involved in interpretation of
the depositional events and stratigraphy of cave sediments, particularly noting the impor-
tance of recognising lateral facies change, secondary unconformities and reverse stratig-
raphy. A deposit such as the floor of Cathedral Cave is likely to be the result of a com-
plex sequence of events over the period of deposition and it is to be expected that a sec-
tion taken through the deposit, such as the section revealed by the present excavation,
would not necessarily represent a simple depositional time line. With these considera-
tions in mind, this attempt at interpretation of the stratigraphy and time events of this
deposit has combined information from the observed sedimentary sequence with inter-
pretation of other inclusions and the taphonomy of the bones to derive a hypothetical
correlation of events. Radiocarbon dates indicate a depositional time span from approxi-
mately 35,000 BP to about 2,000 BP (in fact up to the present, since the top meter of the
deposit had been too disturbed for analysis).
Taphonomic factors were of prime importance in determining the range of fauna
present in the sediments. Fig. 3 indicates that in most spits of the deposit over 90% of the
bones represent small mammals (<1.5 kg body weight) suggesting the main source of
bone was the remains of prey of owls and, in the case of Phase 1, the Ghost Bat. The
general absence of large bone (Table | indicates the largest bone fragment, from Phase 2,
is approx 23 cm long) suggests that at no time was there a large entrance to the cave.
Where large species are present they are represented by fragmentary remains of juve-
niles, suggesting that pitfall situations only allowed entry of small animals, or previously
broken remains of larger animals. These observations could well account for the general
absence of many typical late Pleistocene megafaunal species, e.g. Diprotodon,
Zygomaturus, Palorchestes, Phascolonus, which are present in deposits of approximately
the same age from nearby Cuddie Springs (Dodson et al. 1993).
Of the small mammal taxa which have been identified to species level, most could
have inhabited the area at the time of first European settlement, although many would
have been at the extreme of their known distributions (Strahan 1983). Further analysis of
the rodents is currently being undertaken by one of us, M. L. Augee. This group is the
most abundantly represented in the fauna and preliminary analysis indicates that consid-
erable species diversity is represented. It is hoped that the rodent fauna may be of greater
value in reflecting climatic or vegetation differences over the depositional period.
While differences between the three Phases primarily reflect taphonomic factors
rather than climate or ecology, some trends are apparent.
In Phase 1 the presence of the Ghost Bat and the high level of species diversity
indicate a warm climate with complex vegetation communities and a high level of pro-
ductivity. Although the Ghost Bat is now confined to sub-tropical and monsoonal regions
of Northern Australia, its fossil record indicates that it could tolerate more temperate cli-
mates (Molnar et al. 1984). However, it is suggested that they require breeding caves in
which the mean temperature does not drop below 20 degrees C (Nelson 1989). With the
exception of a record of two specimens from Cliefden Caves, 30 km south of Wellington
(Molnar et al. 1984) this record from Cathedral Cave represents the most southern occur-
rence of the Ghost Bat in Eastern Australia. Owls may also have contributed to the small
mammal fauna of Phase 1, and the different contributions from these two volant carni-
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 1S
vores could account for some of the strange shifts in relative abundance of dasyurid and
rodent taxa in the lower spits of the deposit (Fig. 2). Although Ghost Bat remains are
present throughout the lower 1.5 metres of the deposit they are only abundant in a 50 cm
zone of Spits 51-54. Dramatic differences in the proportion of small and larger rodents
and dasyurids in these and adjacent spits, and the peak in occurrence of Acrobates sp. in
Spits 46-54 may reflect the different feeding preferences of the Ghost Bat compared
with owls, which moval became the prime contributors to the fauna after the departure
of the bats. Associated C!* dates suggest that the period of Ghost Bat occupation of the
caves ceased prior to approximately 21,000 BP, and that the peak occupations occurred
prior to 30,000 BP. These data suggest that cooling climate due to the approaching
glacial period may have forced the Ghost Bats to depart.
Among the larger taxa present in Phase 1, Sthenurus oreas and Macropus
(Notamacropus) n. sp. represent Pleistocene species which do not occur later in Phase 2.
Phascolarctos cinereus, Sarcophilus harrisii, Phascogale calura and unidentified species
of Bettongia, Onychogalea and Thylogale are all confined to Phase 1, supporting the sug-
gestion of complex ecological conditions supporting high productivity during that period.
The presence of a species of land snail characteristic of wet forest habitats also supports
a complex moister environment near the caves during the period represented by Phase 1.
Throughout Phase 2 there is a somewhat higher proportion of large mammals rep-
resented (Fig. 3), and the preservation of bone and nature of the sediments suggests a
very different mode of deposition from that in Phase 1. Larger species are represented
almost entirely by juveniles (Table 3) and by small highly fragmented remains. Extinct
Pleistocene species are present, but rare, the grazing macropodines being the most com-
mon, with notable absence of browsing taxa (except for one jaw fragment of
Protemnodon sp. in Spits 29-33). It is difficult to find firm evidence for climatic change
during this period since the fauna continues to suggest a complex, relatively productive
environment with many microhabitats being sampled. The fauna does not suggest that
the Wellington area suffered undue climatic stress during the glacial maximum, the peri-
od most likely represented by Phase 2 of the deposit. The absence of Cercartetus sp. and
of Burramys parvus, both of which are commonly present in faunas of similar age from
nearby eastern highland regions (e.g. Jenolan Caves, Wombeyan Caves) suggest that the
climate was not unduly cold.
While most of the taxa of small mammals from Phases | and 2 are still extant, and
could have inhabited the Wellington area at the time of first settlement, the macropodine
taxa, in particular, indicate significant change since late Pleistocene times. The smaller
potoroid and macropodine taxa, e.g. Aepyprymnus rufescens, Bettongia sp., Petrogale sp,
Onychogale sp., and possibly Potorous sp. and Thylogale sp. all now inhabit contracted
ranges in the modern fauna but may have been represented in the Wellington area last
century, although they were probably always rare (Strahan 1983, Dickman 1993, 1994).
However, the larger wallabies and kangaroos (Macropus spp.) of the Cathedral Cave
fauna are completely different from the species inhabiting the area today (i.e. M. rufo-
griseus, M. robustus, M. giganteus and Wallabia bicolor). M. giganteus is represented by
isolated teeth in Phase 3 of the deposit (Holocene age), but only its ‘giant’ precursor, M.
(M.) titan is present in Phase 1 and Phase 2. Similarly the Pleistocene precursor of the
Euro, M. (Osphranter) altus is represented in Phase | and Phase 2.
Of the three species of M. (Notamacropus) (‘wallabies’), present in the deposit,
two, M. cf. M. (N.) agilis, and M. cf. M. (N.) dorsalis, are very close to the species now
confined to northern and northeastern Australia and have not been recorded from further
south in historic times. The third species of M. (Notamacropus) is clearly derived from a
new species to be described from the early Pleistocene Bone Cave fauna of Wellington
Caves (Dawson in prep.), where it forms one of the more abundant elements in that
fauna. The Bone Cave species is similar to M. (N.) dorsalis, but is much larger and dif-
fers in other significant features.
Proc. LINN. Soc. N.S.W., 117. 1997
76 LATE QUATERNARY SEDIMENTS
The presence of M. (M.) ferragus in the upper part of Phase 2 is of interest, since
this taxon appears to have been relatively common in the late Pleistocene of far western
New South Wales (Marshall 1973, Merrilees 1973) and of the Pleistocene deposits of the
eastern Darling Downs, Queensland (Bartholomai 1975). Its presence in Phase 2 may be
an indication of more arid conditions and open grassland spreading eastwards into the
Wellington area during the glacial maximum. Overall, the species of Macropus in the
Cathedral Cave deposit represent a suite of taxa allied most closely with the fauna of
northern Australian grasslands, rather than with elements more characteristic of
Pleistocene faunas of Victoria or South Australia, or the macropodine fauna of the
Wellington region today.
The top 2.5 to 3.0 meters of the deposit (Phase 3) evidently represents a period
when the cave had a very restricted opening to the surface, insufficient to allow the entry
of large bone. There is no evidence that any animals fell into the cave, nor of scavenger
species. The faunal remains consist of small bones and teeth of small taxa, predominant-
ly rodents, with some peramelids and dasyurids, probably accumulated by owls roosting
in overhangs or crevices near the entrance. The bones may have washed into the cave
with surface soil and debris during periods of rain, or been blown in during high winds.
Although imprecise, associated radiocarbon dates indicate that some elements of
the Pleistocene “‘megafauna’ survived in the Wellington area at least until the glacial
maximum (about 17,000 BP) and possibly later, but not into the Holocene. However, this
representation is very depleted compared with the old collections from Wellington
Caves, most of which are derived from the Mitchell Cave, Bone Cave and the passages
of the phosphate mines (Dawson 1985). This observation is supported by new collections
from the Bone Cave, yet to be described, which contain a much more diverse array of
taxa than described here from Cathedral Cave, and thus support the hypothesis that the
sediments in each part of the caves complex at Wellington represent different ages.
ACKNOWLEDGMENTS
The contribution of many people during the four years when the excavation was underway is gratefully
acknowledged. In particular we are grateful to the Wellington Shire Council for permission to dig in the floor of
the main tourist cave of the complex, and to Mr Arnold Worboys, the Wellington tourist officer at the time, for
his enthusiastic support. Others present at the initiation of the project were Professor Mike Archer and Dr Ken
Aplin of the University of New South Wales, and Dr Alex Ritchie, Australian Museum. Many students from the
University of New South Wales, and several staff and other volunteers assisted with the dig and with sorting of
bones. In particular the contribution of Bill Symons, Chris Brydon, Henk Godthelp and Armstrong Osborne is
gratefully acknowledged. Dr Steve van Dyke of the Queensland Museum assisted with identification of the small
dasyurids, and Mr Steve Blandford and Ms Giselle Hodge, honours students in Zoology, University of New
South Wales, contributed to the interpretation of the rodents and bandicoots, respectively. From 1984 to 1986 the
work was supported by a grant from the Australian Research Grants Scheme to M. L. Augee and Lyndall
Dawson.
Proc. LINN. SOC. N.S.W., 117. 1997
L. DAWSON AND M.L. AUGEE 77
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Taphonomy and Palaeoenvironmental
Interpretation of a Late Holocene Deposit
from Black’s Point Sinkhole, Venus Bay, S.A.
M.C. MCDOWELL
(Communicated by M.L. Augee)
School of Biological Sciences, The Flinders University of South Australia,
Box 2100, Adelaide, SA, 5001.
McDowell, M.C. (1997). Taphonomy and palaeoenvironmental interpretation of a late
Holocene deposit from Black’s Point Sinkhole, Venus Bay, S.A. Proceedings of the
Linnean Society of New South Wales 117: 79-96
The deposit from Black’s Point Sinkhole, Venus Bay Conservation Park, SA, repre-
sents a continuous 3500 year palaeontological and geological record for the late Holocene.
Taphonomic analysis established the sinkhole as a pitfall trap. Palaeoenvironmental settings
were deduced by analysing sediments and fauna. Age was assessed using carbon dating.
Evidence suggests that around 4000BP precipitation was greater than present and the environ-
ment was dominated by closed canopy forests with an understorey and nearby mud flats.
During this period fauna including Isoodon obesulus and Bettongia penicillata accumulated.
From approximately 4000BP to 1O00BP the climate became warmer, drier and more variable.
During this period sea level retreated, forests became more open and the understorey was
greatly reduced. Species including Perameles bougainville, Pseudomys bolami, Sminthopsis
dolichura, Sminthopsis hirtipes and Thylacinus cynocephalus appeared and/or became domi-
nant. A carbon date associated with a T. cynocephalus tooth suggests an age of 3030+60BP
making it the youngest mainland occurrence recorded. Around 1000BP precipitation
increased and climate became slightly less variable, resulting in an increase in forest and
understorey density. Macrotis lagotis appeared and I. obesulus returned while P. bougainville
and many arid zone species retreated. These species changes can be associated with the
increase in density of forest and understorey during the period of increased precipitation.
Manuscript received 2 April 1996, accepted for publication 23 October 1996.
KEYWORDS: holocene, taphonomy, palaeontology, pit fall trap, bone accumulation, sinkhole
INTRODUCTION
Late Pleistocene to Holocene fossil deposits provide a record of faunal communi-
ties and, by inference, floral communities that occupied their surrounding area prior to
European settlement (Baynes 1987). The study of such fossil deposits is of vital impor-
tance, as it increases our knowledge and understanding of past communities, and pro-
vides a baseline against which to measure change. The aim of this study was to use the
sedimentological and palaeontological evidence from the Black’s Point sinkhole within
the Venus Bay Conservation Park, Eyre Peninsula, SA to reconstruct the regional history
of environmental change before European settlement.
Processes that affect fossils tend to bias against, rather than for, preservation. The
prima facie assumption therefore, is that all fossil deposits are a biased representation of
the community from which they were drawn. Many authors (e.g. Douglas et al. 1966;
Voorhies 1969; Peterson 1977; Behrensmeyer 1978, 1982, 1991, 1993; Behrensmeyer et
al. 1979; Behrensmeyer and Hill 1980; Behrensmeyer and Kidwell 1985, 1979; King and
Graham 1981; Wakefield 1982; Andrews and Nesbit Evans 1983; Hoffman 1988;
Andrews 1990; Baird 1991; Lyman 1994a, b; Simms 1994) have investigated the biases
Proc. LINN. Soc. N.S.W., 117. 1997
80
LATE HOLOCENE DEPOSIT
135°00'
er Bay S TUDY
AREA
Yy~
EYRE
. Bay
SOUTHERN port uncon
OCEAN
PORT AUGUSTA
PENINSULA
YORKE
& PENINSULA,
=
©
Bs
Si VINCENT
KANGAROO = ISLAND
Figure 1. Location map of Black’s Point sinkhole, Venus Bay.
Proc. LINN. Soc.
N.S.W., 117. 1997
M.C. McDOWELL 81
affecting bone assemblages and how they can be recognised. Biasing processes include
environmental and substrate conditions proximal to a death site, the mode of accumula-
tion, hydraulic transport, weathering, bioturbation, animal size, animal lifespan, time
averaging (processes such as erosion and redeposition by which fossils of different age
are concentrated so that they appear to be contemporaneous) and excavation practices.
This study investigates the taphonomic biases present during accumulation of the Black’s
Point sinkhole deposit by analysing samples of the fossils and sediments.
This study coincides with a Department of Environment and Natural Resources
(DENR) program to reintroduce the brush-tailed Bettong (Bettongia penicillata) to the
park. It is of interest to DENR to determine what other locally extinct species might also
be reintroduced.
Location
Black’s Point Sinkhole is located at latitude 33°(10°95” S and longitude 134°(26’°84”
E on an island within Venus Bay Conservation Park (established in 1976). Venus Bay is on
the upper west coast of Eyre Peninsula near the eastern edge of the Great Australian Bight
(Fig. 1). The island lies inside Venus Bay proper and has been connected to the mainland in
geologically recent time by a tombolo. It is not named on maps, but is locally known as
Black’s Point, hence the sinkhole’s name.
Black’s Point sinkhole is approximately 4m deep and somewhat bell shaped (Fig.
2). Its non-parallel walls are about 1.2m apart at the floor but narrow rapidly near the
surface to an approximately ovoid entrance measuring 0.6m by 0.9m. The entrance to
Black’s Point sinkhole lies approximately 6m above mean sea level and 53m from the
sheltered bay on a gentle north-westerly slope of approximately 8 degrees. The area is
surrounded by water but protected from the open ocean by the 50-100 metre high lime-
stone/calcaranite cliffs of Cape Weyland (Fig. 1).
Black’s Point sinkhole has an interesting history. Some time ago a well was dug
inside the sinkhole, presumably by a farmer searching for fresh water. The sediment
removed was left in a spoil heap near its entrance. The sinkhole entrance was later
rediscovered by Conservation Park biologists who covered it with wire mesh in the
belief that it posed a danger to the brush-tailed bettongs being released in the park. In
the process they noted fossils in the spoil heaps and realised the potential of deducing
the park’s original (pre-European) fauna. A large proportion of sediment had been
removed from the sinkhole by the well sinking operation, but sediment varying in thick-
ness from 100 to 790 mm, was left in situ around the walls (Fig. 2) from which samples
could be taken.
METHODS
Excavation
The strata within Black’s Point sinkhole were divided into seven 20cm units and
one 30cm unit using the lip of the sinkhole mouth as a datum. Excavation was confined
to a 0.5m wide column of sediment from the NEE sinkhole wall that was chosen for its
high fossil content. Excavation levels were defined with steel pegs and twine using a
spirit level and tape measure to ensure accuracy. The eight levels were then labelled A
through to H from top to bottom (Fig. 2). Fossiliferous sediment was excavated using
dental picks, trowels, soft brushes and dust pans. A large tray was inserted at the base of
the level being excavated to prevent contamination of lower levels. Sediment was lifted
to the surface by hand using a 10kg bucket (as an arbitrary unit measure) which were
emptied into labelled polythene bags. The fossiliferous sediment was wet sieved by Mr
G. Medlin using nested sieves of 5mm, 2.5mm and 1.25mm mesh. Rocks were removed
Proc. LINN. Soc. N.S.w., 117. 1997
82 LATE HOLOCENE DEPOSIT
ae
EtG See as en eee
Sad Gs ES GSD SSS Se Oa a GD NNE WALL
.
444
Vu NY
44 04st
VN NSN
x
444
SCALE: 1: 25
Metres 0 0.5 1.0 Metres
- Chocolate brown clays
- Red brown clays
- Bridgewater Limestone
Figure 2. A cross sectional view of Black’s Point
sinkhole morphology and infilling sediments with
excavation levels indicated.
Proc. LINN. SOc. N.S.W., 117. 1997
M.C. McDOWELL 83
and fossils were allowed to dry. All skeletal elements were sorted and collected from the
remaining sediment and rock using a small brush and forceps, then stored in labelled
vials for identification and use in quantitative analysis.
Faunal Analysis
Small mammal fossil remains were viewed using a binocular microscope following
the procedure described in Andrews (1990). Specimens were identified by comparison
with published data and reference material accessed from The Flinders University of
South Australia (FUSA), the South Australian Museum (SAM) and the collection of Dr
M Smith. Once identified, specimens were allocated reference numbers and stored in
vials labelled with the species name, level of origin, source and reference number. They
have now been lodged with the SAM mammal collection.
Taphonomic Analysis
Using the methods of Andrews (1990), the numbers of skeletal elements per level
(Ni) were counted and their relative abundance (Ri) calculated, based on comparison
with the minimum numbers of individuals (MNI) multiplied by the expected numbers of
each element (Ei). MNI is calculated by counting the most abundant skeletal element
present and dividing it by the number of those elements present in a complete skeleton
and Ei is calculated by multiplying MNI by the number of each element present in a
complete skeleton.
Ni
Ryo eee
* ~ MNI(Ei)
x 100%
Proportions of elements are indicated by four indices: (a) post-crania in relation to
crania were shown by an index comparing five post-cranial elements (humerus, radius,
ulna, femur and tibia) to numbers of mandibles, maxillae and isolated molars; (b) a sec-
ond index compares numbers of humeri and femora to mandibles and maxillae; (c) loss
of distal limbs is shown by an index comparing numbers of tibiae and radii with numbers
of femora and humeri; (d) relative proportions of isolated teeth were calculated with ref-
erence to numbers of empty alveolar spaces in the jaw such that proportions of more than
100% indicate relative loss of jaws and less than 100% relative loss of teeth; this index
was calculated separately for incisors and molars.
Breakage of post-cranial elements was based on separation of each into complete,
proximal, shaft and distal segments, with the proportions of each category calculated
against the whole.
Breakage of skulls was indicated by proportions of complete skulls, proportions of
maxillae retaining the zygomatic process, and molars and incisors lost from the maxilla.
Breakage of mandibles was indicated by proportions of complete half-mandibles
(with no distinction made between left and right), proportions of mandibles with inferior
borders broken, and molars and incisors lost from the mandible.
The age frequency distribution of small mammals was determined by attributing
each cranial specimen to one of five age classes in accordance with the wear observed on
teeth. Age classes are defined as: (1) very young, no wear on molars, (2) young, small
amount of molar cusp wear, (3) middle aged, molar cusps worn near smooth with dentine
clearly visible, (4) old, molar cusps almost worn away completely with only strips of
enamel remaining, (5) very old, molars very worn with only a rim of enamel around a
basin of dentine remaining. A negligible number of broken teeth and no evidence of ele-
ment digestion was observed so these categories of information were not investigated.
Proc. LINN. Soc. N.S.W., 117. 1997
84 LATE HOLOCENE DEPOSIT
TABLE |
Minimum Numbers of Individuals (MNI) for Mammals from different levels in Black’s Point Sinkhole, Venus Bay.
Stratigraphic Level A B C D E
Sample Size (x10kg) 1 4 4 4 1
Muridae Indet.* 9 232 111 332 18
Rattus sp. D, 12 8
Rattus fuscipes 4 ZZ, 93 71
Pseudomys sp. 2 28 73 35
Pseudomys australis 21 2) 5
Pseudomys shortridgei 9 16 3
Pseudomys gouldii l 2 1
Pseudomys bolami 2
Pseudomys occidentalis 1
Notomys sp. cf.. N. mitchelli 2 1
Total Muridae MNI 17 232 200 332 26
Cercartetus sp. cf. C. concinnus 2
Total Phalangeroidea MNI 2
Peramelidae indet. 2 3 2 17
Isoodon obesulus 2 18
Perameles bougainville 4 9
Total Peramelidae MNI 4 7 11 35 0
Macrotis lagotis 1
Total Thylacomyidae MNI 1 0) 0) 0 0
Bettongia indet. 1 3
Bettongia penicillata 3 6 3 7
Macropus eugenii ] 4 8
Total Macropodoidea MNI 5 13 1] 1 0
Sminthopsis sp. 5 1 3 1
Sminthopsis sp. cf. S hirtipes 1 2
Sminthopsis psammophila 1
Sminthopsis sp. cf. S. dolichura 6 1
Parantechinus apicalis ]
Phascogale tapoatafa
Dasyurus sp. cf.. D. geoffroii | ]
Thylacinus cynocephalus |
Total Dasyuroidae MNI 0 13 8 6
Total N° of Species 6 13 16 10 3
TOTAL MNI 33 265 232 377 27
“ Muridae Indet. MNI is based on individual insisors which often represents the total maximum murid MNI.
Proc. LINN. SOc. N.S.W., 117. 1997
M.C. McDOWELL 85
Sedimentological Analysis
Sediment from each surveyed level of Black’s Point sinkhole was petrographically
analysed in thin section to determine differences in composition grain size and grain
shape in attempt to identify the sediment’s source of origin.
Carbon Dating
All charcoal fragments were collected for each level to use in carbon dating. A
series of five carbon dates were obtained for levels A, C, D, E and G from the
Quaternary Dating Research Centre at the Australian National University (ANU).
Several small fragments of charcoal were collected and added together for each level to
make a sample large enough for dating.
RESULTS
Faunal Analysis
Table 1 presents the mammal fauna and the total number of species identified from
each level of Black’s Point sinkhole. It also includes the MNI calculated for each species,
the MNI calculated for each genus and the total MNI for each level. The discovery of a
single thylacine first upper right molar (M‘/) and two mandibles from the dibbler
(Parantechinus apicalis) are of particular interest.
60
50
HB Level A No individuals
4
2 : B§ Level B No individuals
3 fl] Level C No individuals
5 a Level D No individuals
S
is [] Level E No individuals
Zz
eS
20
10
0
AGE CLASS
Figure 3. Small mammal age frequency distribution where category 1 = juvenile, 2 = subadult, 3= adult, 4 =
old, and 5 = very old.
Proc. LINN. Soc. N.S.W., 117. 1997
86 LATE HOLOCENE DEPOSIT
TABLE 2
The quantity (No) and relative abundances (Ri) of small mammal skeletal elements recovered from different
levels of Black’s Point sinkhole.
Level A B C D E
Sample Size (x10kg) 1 4 4 4 1
Skeletal Elements NO Ri NO Ri NO Ri No Ri NO Ri
Mandibles 12 31.60 401 75.70 418 90.10 390 51.80 11 28.90
Maxillae 14 36.80 345 65.10 336 72.40 207 27.50 13 34.20
Incisors Dil 35-50 312 29.40 315 33.90 1221 81.00 61 80.30
Molars 76 33.30 577 18.10 375. 13.50 821 18.10 112 49.10
Femora 12 31.60 152 28.70 262 56.50 125 16.60 6 15.80
Tibia 16 42.10 93 17.50 251 54.10 166 22.00 34 89.50
Pelvi 24 63.20 134 25.30 194 41.80 92 12.20 8 21.10
Calcanea 12 31.60 19 3.40 13 2.80 41 5.40 11 28.90
Astragali 7 18.40 23 4.30 132.80 I thesk0) Sw l3s220)
Humeri 7 18.40 148 27.90 250 46.30 168 22.30 12 31.60
Radii 5 13.20 8 1.50 26 5.60 10 1.30 Ay. Sso3\0)
Ulnae 16 42.10 91 17.20 68 14.70 110 14.60 5 13.20
Scapulae 14 36.80 97 18.30 36 = 7.80 28 3.70 eS.)
Ribs SEZ 0 293 5.00 199 3.60 373 4.10 5) 120
Vertebrae 154 20.30 258 =.2.40 102. 1.10 82. 0.50 51 2.00
Mean Ri 31.07 22.65 29.80 18.89 DY SY
TABLE 3
Indices indicating proportions of skeletal elements.
Stratigraphic Level A B C D E
Index
% post crania/crania 87.8 65.2 122.7 65.5 69.4
%fem.+hum./man.+max 73.1 40.2 67.9 45.4 75.0
% tib.t+rad./fem.+hum. 110.5 47.0 55.1 68.1 194.4
% isolated molars 166.0 40.1 223 64.3 183.6
% isolated incisors 135
Taphonomic Analysis
The results for the taphonomic analysis of Black’s Point sinkhole fauna are shown
in Tables 2—5 and Figs 2 and 3. Levels F, G and H were not included due to small yields
and sample sizes. Mammal fossils appear most abundant in the upper levels but are com-
mon throughout the sinkhole strata. Level E was the lowest unit analysed and contains
mostly isolated teeth and tibia (Table 2). The proportion of post-crania to crania is mod-
erate (Table 3), while breakage for both cranial and post-cranial elements is high (Tables
Proc. LINN. SOC. N.S.W., 117. 1997
M.C. McDOWELL 87
4 and 5). The proportion of isolated teeth suggest a relative loss of jaws. No elements
show the affects of digestive dissolution, polishing or rounding, but many show the
effects of mild chemical weathering and some root dissolution.
Fossils appear better preserved in level D than level E, but proportions of post-cra-
nia to crania are very similar (Table 3). Mandibles and maxillae are relatively more abun-
dant (Table 2) but the degree of both cranial and post-cranial breakage is similar (Table
4). The proportion of isolated molars is less than 100% suggesting a relative loss of teeth,
but the proportion of isolated incisors is much greater than 100% suggesting a relative
loss of jaws (Table 3).
TABLE 4
Type and extent of breakages in small mammal long bones extracted from different levels of Black’s Point
sinkhole.
Stratigraphic Level A B C D E
Breakage No. % No. % No. % No. % No. %
Humeri
Complete Ih IAs) 81 54.6 131 52.4 19 14.1 0 O
Proximal 4 57.1 63 42.6 109 43.6 110 81.5 OHS
Shaft 1 14.3 1.4 6 2.4 Sense BrZ5
Distal 15 143, 1.4 4 1.6 I (O87 0 O
Ulnae
Complete Py MAS 19 20.9 10 14.7 lee OLS) 0 O
Proximal 14 87.5 Te Sal 58 85.3 108 98.2 5 100
Shaft 0 0 0 O 0 O 1 0.9 0 0
Distal 0 O 0 O 0 0 0 O 0 O
Femora
Complete 9 75 61 39.9 129 49.3 21 16.4 0 O
Proximal gy 25) 82 53.6 124 47.3 100 78.1 5 83.3
Shaft 0 0 0 O Bh. Dass) Ie Gy 7)
Distal 0 5.9 QB Bie Bh 0 O
Tibia
Complete 3») Bil) PUN NSad 3, Wau 10 6.0 0 O
Proximal S) Ol) Byh - Sis)o6) 121 48.0 56 33.7 8 23.5
Shaft SesilR25 7a Wied) 53 21.0 72 43.4 PPS). Taio)
Distal 75) 48 31.3 41 163 28 16.9 BO
Radii
Complete 5 100 8 100 26 83.9 10 55.6 2 100
Proximal 0 0 O il 3 3) 16:7 0
Shaft 0 0 0 0 O 0
Distal 0 0 12.9 S Ais 0
Proc. LINN. Soc. N.S.W., 117. 1997
88 LATE HOLOCENE DEPOSIT
TABLE 5
Small mammal Cranial Element preservation for different level of Black’s Point sinkhole.
Stratigraphic Level A B C D E
Skull Breakage:
% complete 0 0 0 0 0
% maxillae with zygoma. 3.6 66.4 54.7 38.8 23.1
% maxilla molar loss: 27.4 34.8 45.3 36.9 42.3
% maxilla incisor loss 85.7 99.7 100 100 100
Mandible Breakage:
% complete 8.3 0 0.3 0) 0
% inferior border broken 58.3 38.9 21.4 54.1 100
% mandible molar loss: 33.3 32.8 31.8 35.0 42.4
% mandible incisor loss 66.7 79.8 48.8 60.5 100
Skeletal element relative abundance is higher for level C than for levels D and E
with mandibles showing greater than 90% relative abundance and many other elements
showing greater than 40% relative abundance (Table 2). The proportion of post-cranial to
cranial elements is significantly higher in level C than levels D and E, but other indices
of skeletal element proportions are comparable. The proportion of isolated molars and
incisors suggests a relative loss of teeth (Table 3). The degree of breakage is lower in
level C than in levels D and E (Tables 4 and 5).
The fossils of level B are well preserved and the proportion of isolated molars and
incisors is less than 100% suggesting a relative loss of teeth. The proportion of post-cra-
nial to cranial elements is lower than obtained for level C, but other indices of proximal
and distal post-cranial elements are very similar (Table 3). The degree of breakage is
comparable to that seen in level C (Table 4 and 5).
The fossils of level A, the uppermost level analysed, show high post depositional
weathering and high levels of breakage (Tables 4 and 5). The proportions of isolated
molars and incisors are both greater than 100% indicating a relative loss of jaws. This is
supported by the degree of mandible and maxillae breakage.
The age frequency distribution attained for all levels (Fig. 3) is slightly skewed
toward aged individuals but generally reflects those age group proportions found in a liv-
ing community.
Sediments and Stratigraphy
Two major stratigraphic units occur in Black’s Point sinkhole. The lower unit
which extends from the top of Level D to the floor consists of thick red-brown amor-
phous clays. No evidence of lamination or reworking was observed. The upper unit
which extended from the bottom of Level C to above Level A consists of chocolate
brown silty clay which again showed no sedimentary structure. The two units are separated
by a very sharp contact.
When petrographically analysed level E was observed to be composed of 58% clay
minerals, 32% gypsum, 7% micro-crystalline calcite (micrite) and 3% opaques (charcoal).
Non-clay sized grains were fine, sub angular and moderately to poorly sorted. Level D
contained 55% clay minerals, 35% gypsum, 5% micrite and 5% opaques (charcoal). Non-
Proc. LINN. SOC. N.S.W., 117. 1997
M.C. McDOWELL 89
clay sized grains were fine, sub-rounded and moderately to well sorted. Some 2—3mm
clay nodules and calcite gravel were also observed, as were fossil root casts.
Petrographic analysis showed that level C is composed of 50% clay minerals,
35% gypsum, 5% micrite and 10% opaques (charcoal). Sand sized grains are predomi-
nantly fine and sub-rounded to rounded. Some coarse clay nodules and calcite gravel
also occur. The sand sized sediments of level B are fine grained, rounded to sub-round-
ed and moderately sorted. They are composed of 40% clay minerals, 40% gypsum, 15%
micrite and 5% opaques (charcoal). Sediments from level A consist of 50% clay miner-
als, 35% gypsum, 10% micro-crystalline calcite (micrite) and 5% opaques (charcoal).
Sand sized grains are fine and moderately sorted. Gypsum grains appear sub-angular to
sub-rounded, while all other grains appear well rounded. In all levels gypsum grains are
fragmentary suggesting that they have been transported into the sinkhole and are not
authigenic.
TABLE 6
Summary of Results obtained for Radiocarbon Dating of Charcoal collected from different levels of Black’s
Point sinkhole.
Code Number Date Level of Origin
ANU-9893 1160+60 BP A
ANU-9892 3030+60 BP C
ANU-9891 4440+70 BP D
ANU-9890 4040+140 BP E
ANU-9889 4300+290 BP G
Dating
The radiocarbon dates obtained from ANU can be seen in Table 6. It shows a par-
tial succession in age from the youngest at the highest stratigraphic level dated, to the
oldest in a lower stratigraphic level, but the three lower most dates obtained are not in
sequence. Carbon dates obtained for level D and level G are not significantly different,
but the age of level D is significantly older than level E. This suggests that the lower
Strata may have been reworked, although there are no supporting sedimentary structures.
Alternatively, contamination of charcoal used to obtain the two lower-most dates with
younger material may have occurred. Because several charcoal fragments were used to
obtain a workable sample size, spurious grains that originated from higher levels may
have been included. Charcoal samples from the lower strata are especially prone to this
as their sample sizes were very small.
Contaminated charcoal samples would be subject to an averaging effect that would
significantly alter the age estimate obtained. Only a small quantity of ‘young’ charcoal
would be needed to decrease an age estimate obtained from a contaminated sample of
‘old’ charcoal. Radiocarbon dates indicate that the majority of sediments and fossils sam-
pled from Black’s Point sinkhole have not been reworked. It is therefore assumed that
any change in fauna, or any disharmonious species pairs do not result from the mixing of
non-contemporaneous sediments or time averaging.
Charcoal associated with the thylacine tooth recovered from level C was dated at
3030+60BP suggesting the youngest date of a mainland thylacine.
Proc. LINN. Soc. N.S.W., 117. 1997
90 LATE HOLOCENE DEPOSIT
DISCUSSION
Accumulation of Black’s Point sinkhole fauna
The morphology of Black’s Point sinkhole renders it unsuitable for den use by
either mammalian or avian predators. This is supported by the lack of predator tooth
marks, digestive dissolution, rounding and/or polishing of fossils. Indices used to compare
proportions of skeletal elements (Table 3) show no clear preference in the destruction of
cranial elements compared with post-cranial elements, or proximal post-cranial elements
compared with distal post-cranial elements. This indicates that the deposit was not accu-
mulated by a mammalian or avian predator, both of which preferentially digest and/or
break proportions of their prey skeletons (Andrews and Nesbit Evans 1983, Andrews
1990). The age frequency distribution (Fig. 3) obtained for all levels suggests an accumu-
lative or chance collection mode rather than a selective one expected of predators.
Elements from all three Voorhies groups (Voorhies 1969) are well represented on
all levels and no evidence of abrasive wear was observed. This suggests that fluvial
transport was not the primary mode of fossil collection.
As can be seen in Table 1 the size and species of animals present for all levels is
highly diverse, ranging from large animals (> 20kg) to very small animals (< 20g)
(Strahan 1988). Many of the species listed in Table | are still thought to inhabit Eyre
Peninsula, but it is of particular interest to note the collection of thylacine and dibbler
specimens. The deposit was initially considered too recent to collect megafauna speci-
mens, while the dibbler is a threatened species that has not previously been recorded
from Eyre Peninsula fossil deposits, but is common in Western Australia.
Some of the species present are nocturnal while others are diurnal. Some are preda-
tors while others are herbivores and still others are omnivores. The vast majority of
species present are terrestrial ground dwellers with only a few scansorial and arboreal
species present in small numbers.
This indicates a collection mode that does not significantly bias for body size,
feeding habits or activity schedules, but does bias against species’ habitual dwelling
areas. The evidence obtained suggests that the primary mode of collection for the fossils
excavated from Black’s Point sinkhole was a pitfall trap.
In a pifall trap assemblage one might expect to find more articulated or at least
associated elements within the undisturbed sediment excavated. The lack of associated
skeletons suggests that a degree of secondary taphonomic biasing has occurred. This is
believed to be the result of minor hydraulic sorting and compaction. In the winter months
after soil saturation has occurred, runoff water cascades into the sinkhole, eroding and
transporting the surface sediment a short distance towards the sides of the sinkhole
before percolating away. Element disassociation might also result from the activities of
trapped animals before their death. The moderate to high levels of chemical weathering
observed on all specimens suggests that this water was mildly acidic. This weathering,
combined with the occasional root mark and frequent discrete layers of roof fall material
(Fig. 2) show that the sinkhole entrance was becoming larger with time.
Palaeoenvironmental Interpretation
Pitfall traps are extremely useful for interpreting past environmental conditions
because unlike other coliection modes such as predators or fluviatile systems they are geo-
graphically stationary. One can therefore assume that compositional variations will be the
result of changing environmental conditions rather than changing collection agents.
Assuming uniformitarian principles apply, a record of environmental conditions may be
obtained by examining the preferred conditions of animals that once lived proximal to, and
were trapped in a pitfall. Through analysing the fossils from each level of Black’s Point
sinkhole strata, a story of past environmental conditions and changes begins to emerge.
Proc. LINN. SOc. N.S.W., 117. 1997
M.C. McDOWELL 91
TABLE 7
Preferred habitats of mammals from different levels in Black’s Point Sinkhole, Venus Bay.
Stratigraphic Level A B C D E
Sample Size (x10kg) 1 4 4 4 1
Ubiquitous species
Thylacinus cynocephalus J
Percentage ubiquitous species 0 0 7 0 0
Forest species
Rattus fuscipes J J J A J
Pseudomys occidentalis J
Bettongia penicillata J J J J
Macropus eugenii J J J
Cercartetus sp. cf. C. concinnus 4
Isoodon obesulus J J
Phascogale tapoatafa J
Percentage forest species 66.7 30.8 28.6 40 66.6
Heathland species
Pseudomys shortridgei J J J
Parantechinus apicalis J J
Percentage heathland species 0 7.8 12.5 20 0
Desert species
Pseudomys australis J J J
Pseudomys gouldii J J J
Pseudomys bolami J
Notomys sp. cf.. N. mitchelli J J
Sminthopsis sp. cf. S hirtipes J J
Perameles bougainville J J
Macrotis lagotis J
Sminthopsis psammophila J J
Sminthopsis sp. cf. S. dolichura 4 J J
Dasyurus sp. cf.. D. geoffroii J J
Percentage Desert species 16.7 61.5 50 30 33.3
The fossil faunas and sediments of levels D and E are from the same stratigraphic
unit and are similar in preservation and breakage patterns, thus suggesting that they were
collected under very similar environmental conditions. Level E lacks many of the species
recorded in level D, but this is considered a function of level E’s sample size (see Table
1). R. fuscipes and P. tapoatafa both commonly occupy forests. P. tapoatafa is arboreal,
suggesting the presence of tall trees. P. shortridgei populates heath-land, while I. obesu-
lus is known to occur in both forests and heathland, and like B. penicillata, prefers areas
with a low shrubby ground cover or understorey. P. gouldii inhabits hummocky grass-
land, while P. australis prefers rocky desert and D. geoffroii is considered ubiquitous
(Walton 1988) (Table 7).
Proc. LINN. Soc. N.S.W., 117. 1997
92 LATE HOLOCENE DEPOSIT
Several allopatric species pairs occur in level D (Appendix I). Several tropical to
temperate heath and forest dwelling species coexisted with rocky desert and hummock
grassland species, suggesting the environment was more equitable than at present and
that a greater range of niches were available (Lundelius 1983). The assimilation of these
different ecologies indicate an environment composed of broken temperate forest-wood-
land with a low shrubby understorey, interspersed with grassland-heathland and rocky or
alluvial surfaced patches, that requires a higher level of precipitation than is available
under present conditions. The high proportion of detrital gypsum present in the sedi-
ments of these levels suggests that an evaporitic setting also existed nearby. This may
have taken the form of a sabkha mudflat or a playa lake from which gypsum was eroded
by the wind.
The fauna identified in levels C and B indicate a change in palaeoenvironmental
conditions. D. geoffroii and T. cynocephalus are considered ubiquitous, and are therefore
of little interpretive use, but R. fuscipes, N. mitchelli and P. occidentalis are known to
mainly populate temperate forests and shrublands. M. eugenii, S. dolichura and P.
bougainville have been seen to inhabit similar environments, but are also known to occur
in heathlands where P. shortridgei and P. apicalis both predominate. Walton (1988)
reports that P. bougainville prefers to hunt on stony ridges and sandhills, or plains behind
beaches. P. australis occupies a temperate rocky desert while P. gouldii inhabits hum-
mocky grasslands. S. hirtipes and S. psammophila are also known to inhabit grasslands,
but are common to sandhills and plains. S. hirtipes has also been recorded in woodlands
and on alluvial plains and is adapted for locomotion on sandy surfaces (Walton 1988)
(Table 7). These species suggest a more arid palaeoenvironmental setting. This is sup-
ported by the sharp change in clay mineral type from the gypsiferous red-brown clays to
the gypsiferous chocolate brown clays. The change indicates a drying and erosion of
nearby mudflats previously indicated by detrital gypsum only. Several pre-European
disharmonious species pairs have again been observed at this stratigraphic level
(Appendix I). Allopatric species pairs suggest a more equitable environment than exists
at present (Lundelius 1983). Temperate heath and forest dwelling species are seen to
have coexisted with several sandy and rocky desert dwelling species, hummock grass-
land dwelling species and open heath dwelling species. The faunal assemblage deduced
from level C suggests a more arid variable environment dominated by open woodlands
that lack an under storey and are interspersed with grassland-heathland and rocky or allu-
vial surfaced patches.
The faunal composition of level B is similar to the previous level with the addi-
tion of P. bolami, an arid zone species, and the loss of P. occidentalis, P. apicalis , D.
geoffroii and T. cynocephalus, scrubland/heathland and ubiquitous species (Walton
1988). These species have only been recorded in small numbers so their presence or
absence may be a chance event. Several allopatric species pairs similar to level C exist
on level B (Appendix I). The addition of more arid species supports the hypothesis that
the environment became drier, and that forests become more open, losing its under-
storey while grassland, heathland and sandy or alluvial surface patches become more
prominent.
The fauna of level A suggests another change in palaeoenvironmental conditions
(Table 7). The continued presence of R. fuscipes, B. penicillata and M. eugenii suggests a
patchy temperate coastal forest still exists. The reappearance of /. obesulus may be inter-
preted as a shift in the understorey vegetation from the open community preferred by P.
bougainville to a more closed community with greater ground cover. The presence of a
single Macrotis lagotis individual suggests that a sandy or alluvial plain may still exist
nearby. No disharmonious pairs exist at this stratigraphic level suggesting the environ-
ment may be approaching those characteristics seen in the present. The faunal composi-
tion for this level suggests a shift towards a more dense vegetation community as was
previously inferred for Level D and E.
Proc. LINN. SOc. N.S.W., 117. 1997
M.C. McDOWELL 93
CONCLUSIONS
The evidence compiled from the study of the sedimentology and palaeontology of
Black’s Point sinkhole has enabled a palaeoenvironmental reconstruction spanning the
last 4500 years. It indicates three major periods of accumulation. Prior to 4000BP the
fauna present suggest that the climate for the Venus Bay region was wetter, more temper-
ate and less variable than at present. The vegetation was dominated by forest-woodlands
with a thick understorey, while a mudflat area existed within the bay. Around 4000BP
evidence suggests that effective precipitation or water availability decreased and environ-
mental conditions became drier and more variable. This resulted in a more open wood-
land with a decreased understorey density, and drying and erosion of mudflats. Around
1O00BP it appears the region became slightly wetter and environmental conditions
became less variable. This would allow forests to recover slightly, becoming more closed
with a thicker understorey.
Evidence in the literature relating to sites across Australia lends support the
palaeoenvironmental synthesis proposed from the Black’s Point sinkhole data. Several
authors (Bowler 1981; Kershaw 1983; Bowler and Wasson 1984; Chivas et al. 1985;
COHMAP 1988; Shulmeister 1992; Chivas et al. 1993; Barnett 1994; Williams 1994)
report wetter periods from 7500-4500BP for the Australian region, with drier conditions
occurring between 3500-—2000BP, and a climatic recovery taking place 2000—1000BP.
The community composition of the Black’s Point sinkhole fauna appears to be a
mixture of western and eastern species, but it most closely resembles the Nullarbor Plain
and associated coastal fauna deduced by Baynes (1987). This implies a greater range for
the Nullarbor coastal community, extending it to the eastern side of the Great Australian
Bight.
Taphonomic analysis indicates that Black’s Point sinkhole has operated as a pitfall
trap during the Late Holocene. The fossil fauna identified in the sinkhole is considered a
good representation of the terrestrial palaeocommunity that populated the Venus Bay
coastal regions prior to European settlement.
The Black’s Point sinkhole fauna includes the youngest occurring mainland speci-
men of Thylacinus cynocephalus. It suggests that Thylacine populations that once ranged
across southern Australia began to retreat eastward approximately 3000BP before
becoming extinct on the mainland and later in Tasmania. An Accelerator Mass
Spectrometer carbon date from the tooth enamel would confirm its age.
Further palaeoenvironmental information could be obtained from the study of
pollen and land snail shells in the Black’s Point sinkhole strata. This could test and/or
improve the resolution of the palaeoenvironmental interpretation that has been construct-
ed from the palaeontological and sedimentological data collected. More fossil sites
should also be sought out and investigated in the Eyre Peninsula region to better deter-
mine the regional pre-European faunal composition. This will be of use to DENR in their
attempts to re-establish at least part of Eyre Peninsula’s pre-European fauna within the
Venus Bay Conservation Park.
ACKNOWLEDGMENTS
I would like to firstly acknowledge the aid of Prof R. T. Wells, Prof C. C. von der Borch and Dr M. J.
Smith, all of whom provided critical comment, encouragement, and advise during the work on this project. I
also thank Mr. D. Armstrong for his aid in accessing and excavating the site, and Mr. G. Medlin who intro-
duced me to the study, and gave freely of his time, aiding with excavation, advice, identification of dasyurid
species, and arranged funds, supplied by the Department of Environment and Natural Resources ‘Wildlife
Research Fund’ for the carbon dating of five charcoal samples.
Proc. LINN. Soc. N.S.W., 117. 1997
94 LATE HOLOCENE DEPOSIT
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Proc. LINN. SOC. N.S.W., 117. 1997
M.C. McDOWELL 95
APPENDIX I
Allopatric species present on different levels in Black’s Point Sinkhole, Venus Bay
Species Present Allopatric pairs (Pre-Europeans)
(1) Rattus fuscipes 5 and 14
(2) Pseudomys australis 3,5, 6, 17 and 9
(3) P. shortridgei 2,5, 6, 14 and 15
(4) P. gouldii 5 and 17
(5) P. bolami 1,2, 3,4 and 7
(6) P. occidentalis 2,3, 14 and 15
(7) Notomys mitchelli 5 and14
(8) Cercartetus concinnus
(9) Isoodon obesulus 2 and17
(10) Perameles bougainville
(11) Macrotis lagotis
(12) Bettongia penicillata
(13) Macropus eugenii 14
(14) Sminthopsis hirtipes 1, 3, 6, 7 and 13
(15) S. psammophila 3 and 6
(16) S. dolichura
(17) Parantechinus apicalis 2,4 and 9
(18) Phascogale tapoatafa
(19) Dasyurus geoffroii
(20) Thylacinus cynocephalus
Proc. LINN. Soc. N.S.W., 117. 1997
it) aes,
ny ceil ons _- =
Some Smaller Macropod Fossils of South Australia
J.A. MCNAMARA
(Communicated by M.L. Augee)
South Australian Museum, North Terrace Adelaide SA 5000
McNamara, J.A. (1997). Some smaller macropod fossils of South Australia. Proceedings of
the Linnean Society of New South Wales 117: 97-106
Bones of Lagostrophus fasciatus, Lagorchestes asomatus, L. leporides, Onychogalea
lunata, O. fraenta, are reported. Old records of Caloprymnus campestris and Lagorchestes
conspicillatus are rejected. Juvenile material of L. asomatus is described and Bettongia pusilla
sp. noy. is described from the Nullarbor.
Manuscript received 19 April 1996, accepted for publication 24 July 1996
KEYWORDS: Lagostrophus, Lagorchestes asomatus, Lagorchestes, Onychogalea,
Caloprymnus, Bettongia pusilla.
INTRODUCTION
The bulk of the South Australian Museum’s fossil collections are mammal bones of
late Pleistocene to Holocene age. Since 1988 systematic sorting and data collation of these
vertebrate fossils has yielded interesting observations. The new information generated
relates to, past distributions of extant or recently extinct species, new material of rare forms
and some new forms that have been brought to light, and some of it is reported here.
Sound determinations of smaller macropod taxa appear difficult but repay the
effort. Lagostrophus fasciatus, more readily associated with southern Western Australia,
may be extended with specimens from near Ceduna, the River Murray and the South
East. The modern extinction of Lagorchestes species must be anthropogenic but may
well involve synergistic stressors. For Lagorchestes asomatus the second skull known,
first juvenile material, first South Australian record and a huge extension of range were
discovered among material from the enigmatic, ?late Pleistocene Mairs Cave deposit of
the Flinders Ranges. The recently exterminated Lagorchestes leporides, although poorly
known as a living species, is present in Pleistocene and Holocene deposits in the South
East, River Murray, Yorke Peninsula and Flinders Ranges. Some earlier determinations
of Lagorchestes conspicillatus appear to be in error. Onychogalea lunata is recorded
from the Nullarbor, Flinders Ranges and Yorke Peninsula. Onychogalea fraenata may
now be recorded from Kangaroo Island in an apparently Holocene assemblage of mam-
mals including Lasiorhinus latifrons.
Bettongia pusilla sp. nov. is described from material of Koonalda and Weekes
Caves, Nullarbor Plain. It appears to be the taxon, misidentified as Caloprymnus
campestris, sensu Lundelius and Turnbull and known informally as ‘Thompson’s
unnamed potoroid’.
MATERIALS AND METHODS
All material referred to is part of the fossil collections held by the South Australian
Museum, wherein all registered specimens bear numbers with the prefix P, or part of the
mammal collections where the prefix M is used. Teeth are numbered according to my
Proc. LINN. Soc. N.S.w., 117. 1997
98 SMALLER MACROPOD FOSSILS
whim, favouring a system in which adult cheek teeth of macropods are, P3 M1, M2, M3,
M4, with P2 and dP3 shed earlier in life.
All measurements are in millimetres. All localities are within South Australia,
unless otherwise stated. Published measurements of Finlayson’s holotype of
Lagorchestes asomatus were checked against the specimen so that the fossil material
could be measured in a comparable manner.
SYSTEMATICS
Family MACROPODIDAE Gray, 1821
Bettongia pusilla sp.nov. (Fig. 1) = Caloprymnus campestris, sensu Lundelius and
Turnbull, 1984, non (Gould, 1843)
Holotype
P35450 right dentary, juvenile with P>, dP3, My, and Mp, lacking M3, My and Ij, top of
coronoid process and tip of the angular process.
Type Locality
Koonalda Cave, Nullarbor Plain, South Australia, specifically, spit 6 of trench A of the
Gallus site (see Wright 1971).
Age
Holocene from faunal association, although published dates suggest the deposit may be
near 20,000 years old (Thorne 1971).
Etymology
From the latin referring to its small size.
Referred Specimens
Koonalda Cave, Nullarbor Plain: P35446, right maxilla with p2, dP3, M! and es
P35447 part left dentary with My and M3; P35448, right dentary with Ij, P
crypt and M> unerupted; P35449, left dentary with Ij, and My; P35451 right ie
tary with I,, Py and dP3.
Weekes Cave, Nullarbor Plain: P35442, left dentary fragment with M3.
Old Homestead Cave, Nullarbor Plain, Western Australia: P35443 part left dentary with
P3 loose, M), M9, M3 and Mg; P35444, part left dentary with no teeth; P35445,
right Ij.
Diagnosis
A small bettong with the dentary more lightly built and teeth smaller. The molars are
less bulbous, more straight sided, with the crowns proportionately higher with more loph-
like development than the living species of Bettongia. P> has fewer (3) cuspules and asso-
ciated ridgelets, between the anterior and posterior cusps, than B. penicillata, B. gaimardi
and B. lesueur. The condyle is clavate in dorsal aspect, but sub-ovoid in Caloprymnus and
more or less T-shaped in Hypsiprymnodon, Bettongia, Aepyprymnus, and Potorous. It dif-
fers from Caloprymnus in being less robust, and in having, the ramus less markedly bowed
ventrally, the angular process bent inward more markedly, the molars smaller and less bul-
bous, P> finer with three cuspules and ridgelets rather than just one. B. pusilla compares
with Hypsiprymnodon in dentary size but its, premolars are not flared outward, diastema is
shorter, ramus less bowed ventrally and it lacks the distinct hip on the posterior margin of
the masseteric fossa. While it resembles Potorous platyops in jaw size and tooth size, it
Proc. LINN. SOC. N.S.W., 117. 1997
J.A. MCNAMARA 99
Figure 1. Bettongia pusilla sp. novy., holotype dentary, P35450, in dorsal, lateral and medial view, scale = 1 cm.
Proc. LINN. Soc. N.S.W., 117. 1997
100 SMALLER MACROPOD FOSSILS
does not have the dentary form, with ascending ramus swept back at a low angle, the low-
crowned molars, simpler premolar and primitive I, enamel pattern of the potoroo.
Description
The dentary (Fig. 1), 34.2 mm long, is lightly built with a short diastema of 4.6
mm and a depth under M, of 6.4 mm. In lateral aspect the horizontal ramus appears
straight and forms an angle of 60°(with the ascending ramus. The line of the ventral mar-
gin bows downward to its lowest point below the point between M, and M>. This even
curve is interrupted by a slight eminence at the posterior end of the symphysis. The high-
est, most concave portion of the curve is below the masseteric foramen and just before
the angular process which curves downward, inward and backward (to a point below the
condyle in P35451). The posterior margin of the masseteric fossa is produced laterally
only slightly to form a low hip before rising , in a near parallel of the anterior margin, to
the condyle which is clavate in dorsal view. The coronoid process is swept back (to a
point in P35451). The teeth have lengths and maximum widths of: P5, L2.42, W1.80;
dP3, L2.41, W1.87; Mj, L2.80, W2.38; M5, L3.24, W2.57. Po is a small tooth convex
labially and concave lingually with a single crest formed by anterior and posterior cusps
with 3 interposed cuspules and their associated ridgelets. The deciduous dP3 is similarly
small with a blade-like trigonid and loph-like talonid. Mj is a little smaller than M> but
both have similar form with the crown tops nearly as broad as their bases, their four
cusps forming distinct protolophid and hypolophid. The type specimen has the alveoli of
the remaining molars indicating an M3 similar to My and an My which is smaller with a
distinctly narrower hypolophid (and these may be seen as such in P35443). I, is not
retained in the type but the alveolus indicates a smaller more slender version of the tooth
seen in other bettongs. This tooth removed from juvenile dentary, P35448, has enamel on
the lateral and lower surfaces and extending well down toward the open root. It does not
extend as a ridge back along the dorsal lateral edge as in Potorous. The worn IE Lea),
shows a remarkable extension of enamel down along the ventral edge to thé root tip
which is not closed, indicating an extreme hypsodont condition. I note that this tooth is a
good match for the exposed alveolus of jaw fragment, P35443, allowing that they are
from opposite sides of the animal.
Remarks
I am satisfied that the taxon described and figured well by Lundelius and Turnbull
(1984) under the name Caloprymnus campestris is this taxon and note that they found
‘no overlap in any measurement’ (Lundelius and Turnbull 1984: 29) between their sam-
ple from Madura Cave and those quoted by other authorities. Their extensive descriptive
work may be read in conjunction with this formal description of B.pusilla.
The name Calopymnus campestris should be removed from the mammalian record
of the Nullarbor region until verifiable evidence is produced. Tate (1879) listed Bettongia
campestris with a presumed native name of weelba, as a common species while he did
not list Bettognia lesueur, which was perhaps more likely. Finlayson (1932:150) provides
some evidence that ‘Weelba’ does not support Tate’s claim. In this context three old
specimens, M1705, M1706, M1710, all registered in 1922 with locality given only as SA
and no collector named, are now correctly identified as B. Jesueur, but each is branded B.
campestris, in pencil, across the top of the skull. These indicate the possibility of early
misidentification.
Lundelius (1963) lists C. campetris from Webb’s and Snake Pit Caves but particu-
lar specimens are not identified. Archer (1974) lists C.campestris in the fauna of the
Hampton Tableland but does not allow verification by specifying the material in ques-
tion. My own brief inspection of Western Australian Museum material revealed only
B.pusilla and no C. campestris. True Caloprymnus campestris, such as M3256 and
M3257, is a distinctive form with distinctly bowed dentary with fat Bettongia-like molars
Proc. LINN. SOc. N.S.W., 117. 1997
J.A. MCNAMARA 101
and cruder potoroo-like premolars. P> has one ridgelet and P3 has two, whereas B. pusil-
la has 3 and 3 respectively.
Baynes is familiar with B.pusilla (pers.comm.) and has referred to it as “Thomson’s
unnamed Potoroid’ or ‘Unnamed potoroid’ (Baynes 1987). I consider these are one and the
same. While I recognised this taxon independently in South Australia, this was at a later
date than the original work by Peter Thompson (Baynes 1987) and I wish to encourage the
recognition of true discoverers. I favour the use of names that relate directly to the animal
and are mnemonic, and so I suggest Nullarbor Dwarf Betttong for Bettongia pusilla. 1 can
see no practical purpose served by assigning this form to a new genus. I keenly await the
bold cladist who may place it clearly in a diagrammatic phylogeny of the potoroids.
The ecological description of this species cannot be given at the moment, but the
tantalising possibility exists that well preserved mummies may provide some of this
information. Well known from some Nullarbor caves, mummies can preserve pelage,
soft-tissues and gut contents. I urge the collection and proper deposition of such material.
Lagostrophus fasciatus (Péron, 1807)
Specimens
Henschke’s Quarry Cave, Naracoorte, South East: P31639, right dentary; P31640, left
and right maxillae and matching dentaries; P31641, right dentary; P31642, part left
dentary; P31643 and P31644, part left maxillae; P31645, left, 1;. P31646, left I,
and M}, as loose teeth; and seven unregistered loose teeth. ;
Jimmy’s Well, north of Tintinara, South East: P35460, right dentary.
Overland Corner Quarry, River Murray: P33474, right maxilla.
Albert Brown’s Cave, Rocky Point between Ceduna and Penong, West Coast: P31647,
right dentary; P31648, right dentary, juvenile; P31649; right dentary; P31650, bro-
ken skull, adult.
Charra Plains, near Ceduna, West Coast: P31651, skull and right dentary, juvenile.
Point De Mole, Gascoigne Bay, West Coast: P31653 and P31654, a pair of maxillae
belonging to same skull; P31652, left dentary; P31655 and P31656, right dentaries.
Remarks
To the published archaeological evidence from the lower Murray (Wakefield
1964), an early record (Poole 1979) and one specimen reported by Flannery (1990), there
can be added the hard palaeontological evidence of the existence of this species in South
Australia. Its bones have been recovered from a late Pleistocene deposit (Pledge 1990) in
the South East and Holocene deposits from the South East to West Coast.
This taxon may be overlooked in samples of rabbit-sized macropods. It has not
been recorded from the South East (Wells and Pledge 1983) or from Henschke’s (Pledge
1990). In a large sample from Henschke’s Quarry Cave material, in this size range, mini-
mum numbers of individuals of three species were found; 94 Lagorchestes leporides (left
dentaries), 4 Onychogalea lunata (left dentaries) and 4 Lagostrophus fasciatus (left max-
illae, there were 3 left dentaries). So Lagostrophus comprises near 5 percent of this class
from that locality.
This form is easily recognised by the lower molars, with distinctive L-shaped fore-
link lying proud of the anterior cingulum and upper molars with prominent postlink. The
skull, particularly the dentary and lower incisors, present an unusual Sthenurus-like
facies.
Lagorchestes asomatus Finlayson, 1943
Specimens
All from Mairs Cave, Buckalowie, lower Flinders Ranges: P14513 and P35453,
Proc. LINN. Soc. N.S.W., 117. 1997
102 SMALLER MACROPOD FOSSILS
skulls, juvenile; P35454, part skull, juvenile; P14516 and P35455, right dentaries;
P35452, P35456, P35457, P35458, left dentaries. All specimens are juveniles with the
oldest tooth-stage having the M>5 or M2 newly erupted and P> or P* in occlusion.
Remarks
As this is the first juvenile material reported some further observations follow. The
holotype described and figured by Finlayson (1943) is an adult skull with worn molars
and the new fossil specimens conform closely to it. Some measurements of P14513 fol-
lowed by Finlayson’s measurements of the type, M3710, in brackets, for comparison are:
greatest length, 62.3 (65.8); zygomatic breadth, 41.6 (42.9);nasals greatest breadth, 9.2
(9.9);diastema, 10.5 (7.1); palate length 33.7 (36.7). The usual age-related differences are
noted with the older skull a little longer and higher with greater crest development, deep-
er zygomae and bigger bullae. A discrepancy in diastema length results from the cheek
teeth being forward of the orbit by 6.1 (13.5), and of the masseteric process 9.0 (15.3).
This may be explained by forward migration of the teeth, as predicted by Finlayson, and
evidenced by the dentary of the holotype where My is about 5 mm forward of its original
position and P3 is rotated forward in a manner characteristic of the condition found in
macropodines in which tooth progression occurs without loss of that premolar.
The teeth of P14513 measure: P2 length 4.2, width 2.5; dP3, length 4.3, width 3.5;
M! length 4.2, width 3.8. Compared to the type, the first cheek tooth, P2, is shorter than
P», Finlayson’s P* (6.3mm), tapers more markedly, anteriorly, and the lingual cingulum
is less well developed.
The juvenile skull, P14513, of L. asomatus, resembles that of a similiar aged skull
of L. hirsutus, M3102, apart from the posterior breadth and depth of the former, probably
associated with the greater otic development. A comparison of the teeth of these two
species reveals differences. In the first, P> has two distinct cuspules and associated
ridgelets between the anterior and posterior cusps and is longer than the corresponding
tooth in L. hirsutus, which has just one cuspule and ridgelet. The dP3 is more molari-
form, broader anteriorly with a distinct trigonid basin, whereas in L. hirsutus the trigonid
is blade like. My is similar in form but broader and higher crowned in L. asomatus. Of
the upper teeth, P~ is longer with 2 cuspules and ridgelets compared with one in L. hirsu-
tus, and the lingual cingulum is more developed. dP” is broader and more molariform.
M” is broader and higher crowned, while closely matching in form, that of L. hirsutus.
Initial determination and registration in 1968 as Bettongia, submerged this material
in the South Australian Museum and Mr Merv Anderson’s private collection until rede-
termination by the author in June 1994.
Lagorchestes leporides (Gould, 1841)
Specimens
Hereford Stream Cave, lower South East: P29143, an almost complete skull.
Unnamed Cave L106, lower South East: P29144, P29145, P29146, three near complete
skulls.
Dene Kilsby’s Cave, lower South East: P35596, juvenile skull.
Blanche Cave, Naracoorte, South East: P35597, skull.
Henschke’s Quarry Cave, Naracoorte, South East: P17681, right dentary; P17814, left den-
tary; P17826, left dentary; P17798, left dentary, juvenile; P18658, part left dentary;
P17626, part left maxilla; P35598 right dentary; P35599, left maxilla; P35600,
P35602, P35603, left dentaries; P35604 right dentary; P35605 right dentary; P35606,
left dentary; P35607, right maxilla, adult; P35608, left maxilla, juvenile; P35609,
right dentary; P35610, left dentary; and a large collection of unregistered material.
Victoria Fossil Cave, Naracoorte South East; P20275, near complete skull; P25552, part
skull.
Proc. LINN. SOC. N.S.W., 117. 1997
J.A. MCNAMARA 103
Curramulka Town Cave, central Yorke Peninsula: P12921, cemented and crushed skull in
two parts.
Mairs Cave, Buckalowie, lower Flinders Ranges: P35459, left dentary.
Remarks
Historical records indicate that this species was tolerably abundant in the southern set-
tled areas of South Australia (Strahan 1983) and it is not surprising to find its fossil bones
across a wide range. The name L. cf. conspicillatus has appeared on lists for the South East
(Wells and Pledge 1983) but I have seen no specimens to support this. P17626 was regis-
tered as Lagorchestes cf. conspicillatus in early 1972, and indicates a possible source of con-
fusion. The label was amended to ‘/eporides’ at some time but the register entry was not cor-
rected until this year when the number was assigned to a single specimen, previously several
unnumbered specimens of mixed taxa were together under the one label. Two teeth of
Macropus rufogriseus were removed during a later phase of sorting, probably 1989-90.
Lagorchestes, P17814 registered later in 1972, was not given a specific identification and
was subsequently sorted as one left dentary of L. leporides, bearing the number and one left
dentary of Onychogalea lunata now numbered P35470. Confusion of similar sized taxa is
discussed elsewhere. Uncertainty was not confined to the Henschke’s material, P20275 from
Victoria Cave is still registered as Largochestes? [sic] only. In the absence of supporting evi-
dence, L. conspicillatus should be struck from the fossil record of South Australia.
Onychogalea lunata (Gould, 1840)
Specimens
Henschke’s Quarry Cave, Naracoorte, South East:_P35467, left maxilla; P35468, part
right maxilla; P35473, part left maxilla with p3, M!, P35469, left dentary, juvenile,
P35470, left dentary, juvenile; P35474, part left dentary, and some unregistered
loose teeth.
Curramulka Town Cave, Yorke Peninsula: P35466, right dentary.
Corra Lynn Cave, Curramulka, Yorke Peninsula: P35464, juvenile skull, left dentary and
some associated post-cranial bones.
Baldina Creek, near Burrra: P35465, complete juvenile left dentary with P3 exposed in
crypt.
Dempsey’s Lake, Port Augusta: mostly fragmentary material including, P19470, P19493,
P35484, P35486, P35487; and more complete dentaries, P22488 and P35485.
Mairs Cave, Buckalowie, Southern Flinders Ranges: P35461, right dentary juvenile;
P35462 part right dentary, juvenile.
Koonalda Cave, Nullarbor Plain, (Gallus site): P35476, P35477, P35478, P35479,
P35480, P35482, P35483, dentary fragments; P35481, part maxilla.
Purple Goringe Cave, Nullarbor Plain, Western Australia: P35463, mummified upper
thorax with skull and right dentary.
Remarks
Poorly known as a living animal O. /unata seems to have favoured the drier north-
ern and western regions and this is reflected in the occurrence of its bones from the
Nullarbor to the Flinders Ranges. If we can say anything definite about the ecology of
this species, its presence in Pleistocene deposits of the South East, may indicate that
these samples record, at least in part, a drier episode of regional climate.
This species was overlooked in samples of rabbit-sized macropods, as noted else-
where, or simply misidentified. Although its arched tooth-rows, elegantly flaring molar
crests and lophs and vestigial premolars should distinguish it and the following species.
It was not distinguished in various South Australian Museum samples, and Williams
(1982) tentatively discussed material of this and the following species under the heading
of Macropus eugenii.
Proc. LINN. Soc. N.S.W., 117. 1997
104 SMALLER MACROPOD FOSSILS
Onychogalea fraenata (Gould, 1841)
Specimens
Kelly Hill Caves, Kangaroo Island, P35491, skull; P35492, part skull; P35493, left and
right maxillae; P35494, left maxilla; P35490, left maxilla; P35495, P36496,
P35497, P35498, P35499, P35500, dentaries.
Lake Fowler, Southern Yorke Peninsula: P18918, left maxilla; P18919, part left maxilla;
P18920, right dentary with P3 excavated.
Baldina Creek, near Burra: P35488, right maxilla; P21023, part left maxilla; P21090,
part left dentary; P22458, part left dentary.
Dempsey’s Lake, Port Augusta: P18243, P19426, P19469c, fragments with teeth.
Remarks
Some comments for the preceding species apply here, although O. fraenata still
allows ecological study of populations in a tiny part of its former range.
This report seems to include the first record of the species for Kangaroo Island,
where its remains were mixed with those of Lasiorhinus latifrons, now extinct on the
Island, too, but well entrenched on the mainland where it has been well studied ecologi-
cally.
We have evidence of coincidence of the two Onychogalea species from the lower
Flinders Ranges area, but as usual with fossil deposits this should not be taken as proof
of synchrony and, therefore sympatry.
DISCUSSION
The purpose of this paper is to report the results of sorting and systematic re-
organisation of about half the South Australian Museum fossil mammal bone material.
Reports of fossils are all too often associated with speculative interpretation of biologi-
cal, ecological and climatological implications. I can only indicate the potential there
may be for such information to be gleaned from the Quaternary deposits of South
Australia. If reliable determinations of well curated specimens can be married to sound
Stratigraphies with secure, consistent dates of actual bone material and then considered
together with a deeper ecological understanding (if it can be gained from the remaining
rabbit-sized macropods), this may allow detailed explanation of the Pleistocene and
Holocene climate changes as they affected this continent. However, trying to survive as
a museum palaeontologist in an environment increasingly influenced by adverse finan-
cial and intellectual conditions, one should perhaps be content with the modest and
achievable goal of completing the second half of the sort. One hint of what might be
undetected is provided by specimen M1828, associated with labels suggesting that it
was collected in October 1872 by F.W. Andrews in sandhills near Lake Gairdner — it is
a skull of Setonix brachyurus.
ACKNOWLEDGEMENTS
I wish to express my appreciation to all those who collected bone material and lodged it at the South
Australian Museum, including: Peter F. Aitken, Mery Anderson, Fred W. Aslin, Albert H. Brown, Cave
Exploration Group of S. Aust. Inc., A. Gallus, C. Hales, P. Harper, W.A. Head, Ken Heynes, Jack Howard, J.
MacLucas, J. Miekel, H.W. Moeller, G. Pilkington, N.S. Pledge, Ron Simms, D.J. Taylor, A.E. Vigar, Dom
L.G. Williams and Adam Yates. The gestation of a scientific contribution requires a certain sequence of events,
these people were the vital initiator, without whom there would be no knowledge gain. Unlike the biological
model, timing in science is not so definite and in one case, forty years does not seem over long. My thanks to
Lynette Queale for bringing the Setonix skull to my attention, Robyn Cherrington and Lucy Ramsden for typing
and Trevor Peters for photography.
Proc. LINN. SOC. N.S.W., 117. 1997
J.A. MCNAMARA 105
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Proc. LINN. SOc. N.S.W., 117. 1997
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ai
The Distribution of Pleistocene Vertebrates
on the Eastern Darling Downs, Based on the
Queensland Museum Collections
R.E. MOLNAR! AND CORNELIA KURZ?
(communicated by M.L. Augee)
' Queensland Museum, P.O. Box 3300, South Brisbane, Qld., 4101, and
"Institut fiir Palaontologie, Universitat Bonn, Nussallee 8, 53115 Bonn, Germany
Molnar, R.E. and Kurz, C. (1997). The distribution of Pleistocene vertebrates on the eastern
Darling Downs, based on the Queensland Museum collections. Proceedings of the
Linnean Society of New South Wales 117: 107-134
Pleistocene tetrapods have been collected from the eastern Darling Downs of
Queensland for about a century and a half. A search of the registers and audit of the collec-
tions permits a set of faunal lists to be compiled for specific localities for the first time.
Among the tentative conclusions — tentative because of lack of control for collecting bias in
the past, among other factors — are the following. The eastern Darling Downs seems to have
had a uniform vertebrate fauna. Few taxa are found at many localities, and these uncommon
taxa were widespread and either actually rare when alive or subject to preservational bias.
Sthenurine kangaroos were less common than macropodines. Some taxa, including
monotremes, ninja turtles and lungfish seem to actually have been rare when alive.
Crocodiles seem to have been more common and diverse in the northwestern (Dalby) region
of the eastern Downs. Dromornithid birds, medtsoine snakes and ziphodont crocodilians seem
to have been absent altogether.
Manuscript received 27 March 1996, accepted for publication 18 September 1996.
KEYWORDS: Darling Downs, diprotodontids, distribution, macropodids, Megalania, ninja
turtle, Pleistocene, taphonomy.
INTRODUCTION
Historical background
Pleistocene fossils have long been known from the Darling Downs. Australian
Pleistocene fossils were first brought to the attention of the scientific world by discover-
ies in central New South Wales around 1830. Although the Darling Downs was not set-
tled until 1840, Thomas Mitchell had already collected material from the Downs by 1842
(Owen, 1877:240). From that period to 1870 further fossils were found and are included
in Owen (1877), who also published an extract of a letter from Leichhardt describing the
Downs and the occurrences of fossils there (p. 241, also included in Bennett, 1876).
About 25% of the Australian fossil marsupials in the British Museum noted in Lydekker
(1887), 236 of the 835 entries, come from the Darling Downs. This is a minimal estimate
as some entries for material collected by Bennett are located only as “Queensland” and
since Bennett, and his employees, did much collecting on the Downs (Bennett 1876)
some of this material probably derives from the Downs. In fact, the area of the Downs
that produced fossils as given by Bennett is basically the same as that recognised now,
except that it has been extended southeast to include Freestone Creek and Warwick.
However, Bennett noted that few bones had been found in the Dalby region and nothing
was reported from that area in Lydekker (1887). In spite of this long history, the only
attempt to present an overall picture of the region during the Pleistocene is that of
Bartholomai (1976).
Proc. LINN. Soc. N.S.W., 117. 1997
108 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Geography
The Darling Downs comprises low, rolling hills and plains in southeastern
Queensland bounded by the Great Dividing Range on the northeast. It extends from the
Bunya Mountains (and the Main Range to their south), south to the granite belt at the
border with New South Wales, west to the Herries Range and the hilly country west of
the Condamine and north to the hills of the Dividing Range (French, 1989).
Topographically the land gradually descends from the hills, slopes and valleys of the
Main Range in the east, to the plains of the Condamine in the west. In the early nine-
teenth century the Downs was covered with grass and herbs, with open woodland on the
hills (Leichhardt in Owen, 1877). The fossil-bearing part of the Darling Downs is a
roughly rectangular region about 200 km long by 80 km wide orientated with its long
axis parallel to the Great Dividing Range, that is northwest to southeast (Fig. 1). The
region east of the Condamine is known as the eastern Darling Downs, and provides a
Pleistocene fauna, whilst the area to the west (including Chinchilla) is the western
Downs: the Pliocene Chinchilla fauna is sometimes known as the western Downs fauna.
This study is restricted to the Darling Downs east from Macalister and the
Condamine River, and hence the term ‘Downs’ as used here refers to the eastern Darling
Downs not the Chinchilla region or the trans-Condamine portion, with two exceptions.
The two Pleistocene localities west of the Condamine River (Kupunn and Boiley’s prop-
erty), so technically not on the eastern Downs, are included. The sites discussed are
given in Fig. 1, except for a few that can no longer be located.
Stratigraphy
Woods (1960) provided an overview of the eastern Downs Pleistocene sediments.
They consist largely of dark clays, sands and grits derived from the basalts of the
Dividing Range, although the sands may also derive from weathering of the underlying
Mesozoic beds (Gill, 1978). Calcareous nodules are common and carbonate lenses may
be found. Judging from the discoveries of fossils in wells, the fossiliferous sediments are
at least 50 metres deep in places but, as pointed out by Bartholomai (1976), these are
probably of Pliocene age at that depth. Pleistocene sediments are, however, at least 42
metres thick in the northwest (Bennett, 1872). The fossils generally derive from the dark
clay soils but, especially in the northwest along the Condamine, may also be abundant in
the yellowish quartz sands.
Macintosh (1967) and Gili (1978) gave a stratigraphy of the region between Kings
and Dalrymple Creeks introducing the ‘Toolburra Silt’ and overlying “Talgai Pedoderm’
and ‘Ellinthorpe Clay’, names which have not subsequently seen general use. Gill gave a
chronology of depositional events in the Dalrymple Creek region. The Toolburra silt was
deposited with brief intervals of lower energy (marked by clay deposition) and higher
energy (marked by sand) flows. This deposition was followed by a dry period that oxi-
dised the Toolburra, and produced some carbonate nodules about 26,000 years old. The
Talgai and Ellinthrope were then deposited, and record periods of dryness and flooding
respectively. Gill interpreted the climate as having generally been wetter than at present.
Macintosh suggested that the intermediate and sporadically occurring ‘Talgai Fossil Soil’
was deposited about 12,000 years ago, based on carbonate dates from the Toolburra and
Ellinthorpe. Gill provided dates for Kings Creek sites as old as 40,000 years. These two
workers made the only attempts at dating the Downs fossils or deposits.
Gill discussed the dates based on the carbonate nodules (and Gill reports that most
dating laboratories were reticent about using these), although in his Fig. 2 he indicated
that comparable dates were also obtained from charcoal (Gill, 1978). Some dates were
also derived from bone and shell according to the figure. Terrestrial carbonates are now
regarded as the least reliable material for radiocarbon dating with whole bone and shells
not greatly better, but charcoal is regarded as probably the best (inorganic) material for
Proc. LINN. SOC. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ 109
¢ Chinchilla
79 yimbour CU
Cattle Cr.
9
3
BY 11 ®@ >
palby ® 40 series
13 yall Ck.
@
Cr 24
26 27
Gowrie
* Toowoomba
28 @
44 42 45 Spring o,
46
Warwick
Gore
\ Brisbane
Figure 1. Pleistocene fossil localities of the eastern Darling Downs. The diamonds represent selected major
towns (four of which are also localities) and the dots (other) localities. The Condamine River and major creeks
are also shown. By region, the localities are: Dalby region: 1, Jimbour; 2, Brimblecomb’s and Jimbour Ck, c.
1.5 km S of Jimbour; 3, Pirrinuan and Jimbour Ck. c. 4.5 km S of Jimbour; 4, ‘Wyoming’; 5, c. 3 km down-
stream from ‘Armour’ and ‘Darrington’; 6, Macalister; 7, ‘The Myalls’; 8, Boiley’s; 9, Kaimkillenbun; 10,
“Crystal Brook’; 11, Mocatta’s Corner; 12, Kapunn; 13, Loudon’s bridge; 14, ‘Territ’; and Dalby. Cecil Plains
region: 15, ‘Cardoch’; 16, ‘Springvale; 17, St. Ruth; 18, Irongate; 19, Bongeen; 20, ‘Cecil Downs’; 21,
Braemar. Toowoomba region: 22, Balgowan; 23, Goombungee; 24, Oakey; 25, near Kingsthorpe (Gowrie
Ck.); 26, Kingsthorpe (Westbrook Ck.); 27, Gowrie; 28, Wellcamp; 29, ‘Eton Vale’; 30, Cambooya; 31,
‘Harrow’; 32, ‘Woodstock’ and ‘Cowarrie’; and 33, Greenmount. Clifton region: 34, Hirstglen; 35,
‘Ravensthorpe’; 36, Brown’s, Bell’s, ‘Greenfields’ and Pilton; 37, ‘Manapouri’; 3, Pearson’s and Budgie
Creek; 39, Nobby; 40, ‘Bundah’; 41, Clifton; 42, Sutton’s; 43, College Green; 44, O’ Mara’s bridge; 45, Spring
Creek; 46, Talgai; 47, ‘Ebley’; 48, 6.5 km W of ‘Goomburra’; 49, ‘Goomburra’; 50, Eastwill’s; 51, Freestone;
52, Yangan. In addition the locality at Gore is also shown. Not all of these localities are known with compara-
ble accuracy, e.g., ‘Ebley’ cannot now be located, other than that it was near Allora, so the dot marks Allora.
Glengallen Plains could not be located. The regions defined in the text are shown by dashed lines.
Proc. LINN. Soc. N.S.W., 117. 1997
110 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
dating (Meltzer and Mead, 1985). However the strength of the association of the charcoal
(and shells and nodules) with the bones is not given. In view of these considerations it is
very desirable to replicate these dates.
Aims of this work
This study was intended to achieve five aims:
1). to provide a general introduction to the Downs and its fossils for paleontologists,
2). to collect together the data in the Queensland Museum on vertebrate fossils from the
eastern Downs,
3). to test regionalism in the region,
4). to check whether there was one, or more, local fauna, and finally,
5). to see if there were any interesting conclusions to be drawn from the previously unin-
spected data.
It has been generally assumed that the distribution of fossil taxa on the eastern
Downs is uniform, i.e. that there is no regionalism. Because of the importance of this
area for the understanding of Pleistocene Australia, it is desirable to check this assump-
tion. This study was undertaken in part in response to the comment of Archer (1984) that
eastern Darling Downs local fauna might actually be more than a single fauna. And
because the fauna is undated, it has also been assumed that the fossils were all more or
less of the same age, although admittedly several workers have emphasised that this may
not be the case. Dating the material and taphonomic observation, such as whether the
sites represent high or low energy deposition, unfortunately cannot be addressed here.
Even so it is clear that some sites, e.g. Sutton’s and Pearson’s, seem to represent low
energy deposition while others, e.g. “~Bundah’, had at least some episodes of high energy
deposition.
Caveats of this work
Several other caveats of this study must be noted. It is restricted to the eastern
Downs and is based only on material (including casts) in the Queensland Museum col-
lections. This is probably the largest collection of eastern Downs material, and so lends
confidence to the conclusions, but still the study is indicative rather than comprehensive.
In view of the large number of specimens involved, by and large the collection identifi-
cations were not verified but simply accepted. The mammalian nomenclature has been
updated following Archer et al. (1984). However, all register entries were checked in a
thorough audit of all specimens not presently on loan.
In view of the lack of taxonomic revision of Pleistocene diprotodontids (now
underway by B. Mackness) these have been put into three categories: identifications as
Diprotodon australe, D. australis and D. optatum are given here as “Diprotodon, large
form”; Diprotodon minor as “Diprotodon, small form’, and all other, non-Diprotodon
diprotodontids as “small diprotodontids” (which probably includes several species).
Likewise crocodilians (almost invariably recorded as Pallimnarchus pollens) have been
given as “crocodilian” except where diagnostic features have been preserved.
Chelonians, unless clearly referable to the Meiolaniidae or the Trionychidae, have been
given as chelids.
Similarly locality identifications have been assumed to be accurate. Some of the
early localities, such as ‘Gowrie’ or ‘Pilton’, probably refer to regions rather than to dis-
coveries at those specific locations. Hence some taxa in the lists for these localities may
actually be referable to other, more specific localities included here such as, e.g.,
Brown’s property at Pilton. Thus some taxa may be included in the wrong lists, or some
Proc. LINN. SOC. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ 111
of the localities given as separate may be identical. For much of the early work this
would seem prohibitively difficult or impossible to determine now, and it is hoped that
this does not greatly alter the conclusions. It should be remembered that these lists are
lists of fossils found at the localities. Faunal lists, of which taxa lived in these regions,
may be compiled from the lists of fossils but we have not done so. Thus discrepancies
between lists for nearby localities reflect differences in the fossils (so far) found, but do
not necessarily imply differences in the kinds of animals that lived there.
Most sites have not been systematically collected, and collecting before 1970 was
probably restricted to collecting vertebrates of moderate to large size, about the size of
Sarcophilus and larger. Recently, systematic collecting has been done at some sites,
specifically Sutton’s, Pearson’s, O’Mara’s Bridge and ‘Bundah’. The last site, ‘Bundah’,
has been exhaustively collected including everything uncovered at the locality since its
discovery.
Because of the lack of control over early collecting and other constraints a detailed
statistical analysis of the faunal lists is unwarranted. Such an analysis would seem to pro-
vide an uncomfortably large possibility of “garbage in, garbage out’. Therefore only a
general set of suggested conclusions — working hypotheses — based on inspection of
the data for the distribution of Pleistocene taxa on the eastern Downs is presented. A list
of the tetrapod material in the British Museum in the late 19th century from the eastern
Downs localities, taken from Lydekker (1887, 1888, 1889) but with the nomenclature
updated, is given in the Appendix |. Since there has not been much collecting on behalf
of the British Museum on Downs since then, this probably gives a reasonably complete
list of their eastern Downs collection.
FOSSIL LISTS FOR EASTERN DARLING DOWNS LOCALITIES
Synonymies from Archer, et al., (1984), Ingram, (1990), Van Tets and Rich (1990).
The number in parentheses before most localities is the locality number in Fig. 1. Note
that nearby localities are not individually represented by dots on the figure, and that
imprecise localities,such as “Jimbour Creek”, and major towns, such as Warwick, are not
numbered. The number in parentheses after the locality name gives the number of taxa
found at that locality.
DALBY Region: 20 localities + “‘Jimbour Creek”
Site 1 — Jimbour (2):
Macropodinae
Macropus ?ferragus
Protemnodon anak
Site 2 — Jimbour Ck., c. 1.5 km S of Jimbour (1):
Diprotodontidae
Diprotodon, large form
Site 2 — Brimblecomb’s property, Jimbour Creek (3):
Macropodinae
Macropus titan
Protemnodon anak
Diprotodontidae
Diprotodon, large form
Site 3 — Jimbour Ck., c. 4.5 km S of Jimbour (1):
Sthenurinae
Procoptodon goliah
Proc. LINN. Soc. N.S.w., 117. 1997
112 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Site 3 — Pirrinuan, Jimbour Creek (1):
Sthenurinae
Troposodon minor
Site 4 — ‘Wyoming’ (2):
Macropodinae
macropod
Diprotodontidae
?Diprotodon, large form
Site 5 — Condamine River, c. 3 km downstream from ‘Armour’ (2):
Diprotodontidae
Diprotodon, large form
small diprotodontid
Site 5 — near ‘Darrington’, Condamine River (1):
Diprotodontidae
Diprotodon, large form
Site 6 — Macalister, Condamine River (16):
Vombatidae
Phascolomys sp.
Vombatus sp.
Macropodinae
Macropus ferragus
Macropus pearsoni
Protemnodon anak
Protemnodon roechus
Sthenurinae
Procoptodon pusio
Troposodon minor
Diprotodontidae
Diprotodon, small form
Diprotodon, large form
small diprotodontid
Palorchestidae
Palorchestes azael
Crocodylidae
Crocodylus porosus
Pallimnarchus pollens
Meiolaniidae
meiolaniid
Chelidae
chelid
Site 7 — ‘The Myalls’ (and vicinity) (3):
Macropodinae
Protemnodon anak (c. 1 km N.)
Diprotodontidae
Diprotodon, \arge form
small diprotodontid
Site 8 — Boiley property, near ‘Daandine’ (1):
Macropodidae
macropod
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R.E. MOLNAR AND C. KURZ
Site 9 — Kaimkillenbun (3):
Vombatidae
Vombatus sp.
Macropodinae
Protemnodon brehus
Sthenurinae
Sthenurus andersoni
Site 10 — ‘Crystal Brook’, Myall Creek (2):
Macropodinae
macropod
Diprotodontidae
diprotodontid
Site 11 — Mocatta’s Corner, Bunya Creek (1):
Varanidae
Megalania prisca
Site 12 — Kupunn (given “Kapunn’’) (1):
Vombatidae
Phascolonus gigas
Site 13 — ‘Greenbank’ (Loudons Bridge) (1):
Diprotodontidae
diprotodontid
Site 14 — ‘Territ’, Oakey Creek (1):
Vombatidae
Phascolomys medius
Dalby region, Cattle Creek (9)
Vombatidae
wombat
Hypsiprymnodontinae
Propleopus oscillans
Macropodinae
Macropus ferragus
Macropus titan
Protemnodon anak
Protemnodon roechus
Diprotodontidae
Diprotodon, large form
Dasyuridae
Sarcophilus laniarius
Thylacoleonidae
Thylacoleo carnifex
Dalby (12):
Vombatidae
Phascolomys magnus
Macropodinae
Macropus titan
Protemnodon anak
Wallabia indra
Proc. LINN. Soc. N.S.W., 117. 1997
114 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Diprotodontidae
Diprotodon, \arge form
small diprotodontid
Palorchestidae
Palorchestes azael
Thylacoleonidae
?Thylacoleo carnifex
Dromaiidae
Dromaius sp. cf. D. novaehollandiae
Crocodylidae
crocodilian
Varanidae
Megalania prisca
Chelidae
chelid
‘‘Jimbour Creek” (6):
Macropodinae
Macropus agilis
Macropus titan
Protemnodon anak
Protemnodon roechus
Diprotodontidae
Diprotodon, large form
Thylacoleonidae
Thylacoleo carnifex
Myall Creek (1):
Macropodidae
macropod
‘CONDAMINE RIVER’
Macropodinae
Macropus ferragus
Diprotodontidae
Diprotodon, large form
small diprotodontid
Dromaiidae
Dromaius sp.
Crocodylidae
crocodilian
Varanidae
Megalania prisca
Chelidae
Chelodina sp. (given “Chelonia’’)
CECIL PLAINS Region: 8 localities
Site 15 — ‘Cardoch’, Condamine River (1):
Macropodinae
Protemnodon anak
Site 16 — ‘Springvale’, near Tipton (5):
Macropodinae
Macropus pearsoni
Protemnodon anak
Proc. LINN. SOC. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ
Diprotodontidae
Diprotodon, small form
small diprotodontid
Palorchestidae
Palorchestes azael
Site 17 — St. Ruth (2):
Diprotodontidae
Diprotodon, large form
Palorchestidae
Palorchestes azael
Site 18 — Irongate (1):
Diprotodontidae
Diprotodon, small form
Site 19 — Bongeen (2):
Macropodinae
Macropus titan
Osphranter altus
Site 20 — ‘Cecil Downs’, Linthorpe Creek(7-8):
Macropodinae
Macropus sp.
Protemnodon roechus
Sthenurinae
Procoptodon sp.
Diprotodontidae
Diprotodon, large form
small diprotodontid
Thylacoleonidae
Thylacoleo sp.
family not known
?bird
Agamidae
agamid*
Site 21 — ‘Braemar’, near Southbrook (1):
Diprotodontidae
Diprotodon, large form
Cecil Plains (1):
Diprotodontidae
Diprotodon, large form
115
* The condition of the specimen suggests that it may be recent or of different age from the
rest of the taxa listed.
TOOWOOMBA Region: 14 localities + ““Gowrie Creek”
Site 22 — Balgowan colliery (2):
Macropodinae
Protemnodon anak
Diprotodontidae
Diprotodon, large form
Site 23 — Goombungee (1):
Macropodinae
Protemnodon anak
Proc. LINN. Soc. N.S.W., 117. 1997
116 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Site 24 — Oakey, Gowrie Creek (1):
Diprotodontidae
Diprotodon, large form
Site 25 — near Kingsthorpe, Gowrie Creek (1):
Macropodidae
macropod
Site 26 — Kingsthorpe, Westbrook Creek (4):
Macropodinae
Macropus titan
Protemnodon anak
Diprotodontidae
Diprotodon, small form
Diprotodon, large form
Site 27 — Gowrie (25):
Vombatidae
Phascolomys angustidens
Phascolonus gigas
Vombatus ursinus
Potoroinae
Aepyprymnus rufescens
Macropodinae
Macropus agilis
Macropus dryas
Macropus thor
Macropus titan
Onychogalea unguifera
Osphranter altus
Protemnodon anak
Protemnodon brehus
Protemnodon devisi
Protemnodon roechus
Wallabia indra
Sthenurinae
Procoptodon pusio
Sthenurus andersoni
Troposodon minor
Diprotodontidae
Diprotodon, large form
small diprotodontid
Thylacinidae
Thylacinus cynocephalus
Thylacoleonidae
Thylacoleo carnifex
Rallidae
Gallinula mortierii
Crocodylidae
crocodilian
Chelidae
chelid
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R.E. MOLNAR AND C. KURZ
Site 28 — Wellcamp and vicinity (5):
Macropodinae
Macropus sp.
Protemnodon anak
Protemnodon roechus
Diprotodontidae
Diprotodon, large form
small diprotodontid
Site 29 — ‘Eton Vale’ (2):
Macropodinae
Macropus sp.
Diprotodontidae
Diprotodon, large form
Site 30 — Cambooya region (3):
Macropodinae
Macropus titan
Protemnodon anak
Diprotodontidae
Diprotodon, large form
Site 31 — ‘Harrow’ (“Sharrow”) (7):
Macropodinae
Macropus pearsoni
Macropus titan
Protemnodon anak
Sthenurinae
Troposodon minor
Diprotodontidae
Diprotodon, large form
small diprotodontid (Prochoerus celer)
Crocodylidae
crocodilian
Site 32 — ‘Cowarrie’, near Southbrook (1):
Diprotodontidae
Diprotodon, large form
Site 32 — ‘Woodstock’, Hodgsons Creek (4):
Diprotodontidae
Diprotodon, large form
family not known
bird
Varanidae
Megalania prisca
family not known
lizard
Site 33 — Greenmount, Emu Creek (6):
Macropodinae
Macropus titan
Protemnodon anak
Sthenurinae
?Procoptodon sp.
117
Proc. LINN. Soc. N.S.W., 117. 1997
118 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Diprotodontidae
Diprotodon, large form
Peramelidae(?)
bandicoot
family not known
bird
‘Gowrie Ck” (9):
Macropodinae
Macropus sp.
Protemnodon anak
Protemnodon roechus
Diprotodontidae
Diprotodon sp.
diprotodontid (Sthenomerus charon)
Dasyuridae
Sarcophilus sp.
Thylacinidae
Thylacinus cynocephalus
Thylacoleonidae
Thylacoleo carnifex
family not known
teleost
Toowoomba (1):
Diprotodontidae
Diprotodon, large form
CLIFTON Region: 25 localities + ‘Kings Creek”
Site 34 — Longe’s property, Hirstglen (1):
Macropodinae
Macropus titan
Site 35 — ‘Ravensthorpe’ (19-20):
Vombatidae
Vombatus sp.
Macropodinae
Macropus agilis
Macropus ferragus
Macropus pan or faunus
Macropus piltonensis
Macropus thor
Macropus titan
Osphranter woodsi
Protemnodon anak
Protemnodon ?devisi
Protemnodon roechus
Sthenurinae
Sthenurus pales?
Troposodon minor
Diprotodontidae
Diprotodon, large form
Rallidae
large rail
Proc. LINN. SOc. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ
Ardeidae
Ardea cf. A. novaehollandiae
Megapodidae
Progura gallinacea
Crocodylidae
crocodilian
Varanidae
Megalania prisca
Trionychidae
trionychid
Site 36 — Bell’s property, Pilton (1):
Palorchestidae
Palorchestes parvus
Site 36 — Brown’s property, Pilton, Kings Ck. (7):
Macropodinae
Macropus agilis
Macropus cf. M. giganteus
Protemnodon anak
Protemnodon roechus
Diprotodontidae
Diprotodon, large form
small diprotodontid
Chelidae
chelid
Site 36 — ‘Greenfields’, Pilton (4):
Vombatidae
Phascolonus gigas
Macropodinae
Macropus ferragus
Macropus sp. cf. M. titan
Protemnodon anak
Site 36 — Pilton, Kings Creek (17):
Vombatidae
Phascolomys angustidens
Phascolonus gigas
Vombatus ursinus
Macropodinae
Macropus ferragus
Macropus titan
Protemnodon anak
Protemnodon roechus
Sthenurinae
Sthenurus andersoni
Diprotodontidae
Diprotodon, large form
Dasyuridae
Dasyurus sp.
Sarcophilus laniarius
Thylacinidae
Thylacinus cynocephalus
119
Proc. LINN. Soc. N.S.W., 117. 1997
120 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Thylacoleonidae
Thylacoleo carnifex
Ornithorhynchidae
Ornithorhynchus anatinus
Dromaiidae
Dromaius novaehollandiae
Varanidae
Megalania prisca
family not known
teleost
Site 37 — ‘Manapouri’, Kings Creek (3):
Macropodinae
Macropus titan
Diprotodontidae
Diprotodon, large form
Chelidae
chelid
Site 38 — Budgie Creek (1):
Macropodinae
macropod
Site 38 — Pearson’s Locality, Kings Creek (22):
Vombatidae
Phascolomys medius
Phascolonus gigas
Macropodinae
Macropus agilis
Macropus ferragus
Macropus pearsoni
Macropus titan
Onychogalea unguifera
Protemnodon anak
Protemnodon brehus
Protemnodon roechus
Thylogale sp.
Sthenurinae
Procoptodon pusio
Procoptodon rapha
Sthenurus andersoni
Troposodon minor
Diprotodontidae
Diprotodon, large form
Palorchestidae
Palorchestes azael
Dasyuridae
Dasyurus sp.
Thylacinidae
Thylacinus cynocephalus
Phasianidae
quail
Varanidae
Megalania prisca
Proc. LINN. SOC. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ
Plotosidae
Tandanus tandanus
Site 39 — Nobby (1):
Rallidae
Gallinula (G. mortierii?)
Site 40 — ‘Bundah’, Neds Gully (9):
Vombatidae
wombat
Macropodinae
Macropus titan
Sthenurinae
Procoptodon goliah
Diprotodontidae
Diprotodon, large form
Dasyuridae
Sarcophilus sp.
Thylacoleonidae
Thylacoleo sp.
family not known
rodent
family not known
?bird
family not known
?teleost
Site 41 — Clifton, Kings Creek (17):
Vombatidae
Lasiorhinus sp.
Phascolomys sp.
Macropodinae
Macropus agilis
Macropus titan
Protemnodon affinis®
Protemnodon anak
Protemnodon devisi
Protemnodon roechus
Osphranter altus
Sthenurinae
Procoptodon goliah
Procoptodon pusio
Troposodon minor
Diprotodontidae
Diprotodon, small form
Diprotodon, large form
small diprotodontid
Thylacoleonidae
Thylacoleo carnifex
Rallidae
Gallinula mortierii
Site 42 — Sutton’s site, Kings Creek (18):
Vombatidae
Phascolonus gigas
Proc. LINN. Soc. N.S.W., 117. 1997
122 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Macropodinae
Macropus titan
Protemnodon roechus
Diprotodontidae
Diprotodon, large form
Palorchestidae
Palorchestes sp.
Peramelidae(?)
?bandicoot
Dasyuridae
Dasyurus sp.
family not known
?monotreme
family not known
rodent
family not known
bird
Crocodylidae
crocodilian
Varanidae
Megalania prisca
other varanid
family not known
snake
Agamidae
agamid
Chelidae
chelid
family not known
?frog
family not known
teleost
Site 43 — near College Green, Kings Creek (1):
Diprotodontidae
diprotodontid
Site 44 — O’Mara’s Bridge, Kings Creek (12):
Macropodinae
Macropus agilis
Protemnodon anak
Sthenurinae
Troposodon kenti
Troposodon minor
Diprotodontidae
Diprotodon, \arge form
Dasyuridae
Dasyurus sp.
Thylacoleonidae
Thylacoleo sp.
Crocodylidae
crocodilian
Proc. LINN. SOC. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ
Varanidae
Megalania prisca
other varanid
Chelidae.
chelid
family not known
teleost
Site 45 — near Clifton, Spring Creek (5):
Macropodinae
Macropus agilis
Macropus ferragus
Macropus titan
Diprotodontidae
diprotodontid
Thylacoleonidae
Thylacoleo carnifex
Site 46 — Talgai (1):°
Macropodinae
Protemnodon anak
Site 47 — ‘Ebley’, near Allora (1):
Diprotodontidae
Diprotodon, large form
Site 48 — 6.5 km W. of ‘Goomburra’ (1):
Vombatidae
Phascolonus sp
Site 49 — ‘Goomburra’, near Allora (1):
Palorchestidae
Palorchestes sp.
Site 50 — Eastwill’s property, Glengallen Creek (1):
Sthenurinae
Procoptodon goliah
Site 51 — Freestone, Freestone Creek (14):
Vombatidae
Phascolonus gigas
Vombatus ursinus
Macropodinae
Macropus agilis
Macropus titan
Osphranter altus
Protemnodon anak
Sthenurinae
Procoptodon goliah
Procoptodon rapha
Sthenurus sp. (nov.?)
Diprotodontidae
Diprotodon, large form
small diprotodontid
Proc. LINN. SOc. N.S.W., 117. 1997
124 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Peramelidae
Perameles sp. (nov.?)
Dasyuridae
Dasyurus sp
Ciconiidae
Ephippiorhynchus asiaticus
Site 52 — Yangan (given “Yangau’’) (1):
Diprotodontidae
Diprotodon, large form
Glengallen Plains (1):
Vombatidae
Lasiorhinus latifrons
“King Creek”:
Vombatidae
Phascolonus sp.
Vombatus sp.
Hypsiprymnodontinae
Propleopus oscillans
Macropodinae
Macropus agilis
Macropus ferragus
Macropus titan
Protemnodon anak
Protemnodon brehus
Protemnodon roechus
Sthenurinae
Procoptodon goliah
Procoptodon rapha
Troposodon minor
Diprotodontidae
Diprotodon, large form
small diprotodontid
Thylacoleonidae
Thylacoleo carnifex
family not known
monotreme
Dromaiidae
Dromaius novaehollandiae
Anatidae
duck
Accipitridae
buteonine (Taphaetus brachialis)
Varanidae
Megalania prisca
“Varanus dirus”
Meiolaniidae
Ninjemys oweni
Chelidae
chelid
Ceratodontidae
Ceratodus palmeri
Proc. LINN. SOC. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ 125
Warwick (2):
Diprotodontidae
small diprotodontid
Columbidae
Phaps sp.
* Locality uncertain.
© The label makes clear that this is a reference to the Macropus affinis of Owen (1845; cf.
Mahoney and Ride, 1975).
“We agree that the human skull wasn’t contemporaneous with Protemnodon (cf. Gill,
1978).
DISCUSSION
Size bias
Almost half of the taxa identified in the lists are recorded from only one locality.
This obscures any geographical patterns that might be present. Such patterns, if any, are
probably not reliable for the small taxa (e.g., birds, Dasyurus, bandicoots, rodents,
monotremes, small lizards, frogs). Collecting experience, especially at Sutton’s site and
‘Bundah’, suggests that these small forms have been overlooked and actually may have
been much more widespread, as would be expected from ecological considerations.
Contrawise any geographical patterns are probably more reliable for the larger
forms so suggested conclusions here are restricted (largely) to these large forms.
Regionalisation
In order to discern any geographical patterns the eastern Downs, west to (and
including) the region around Macalister, was divided somewhat arbitrarily into four por-
tions (Fig. 1): around Dalby; around Cecil Plains; around Oakey and Toowoomba; and
around Clifton and Warwick, respectively called the Dalby, Cecil Plains, Toowoomba,
and Clifton regions. Taxa found in each region are given in Appendix 2. The variety of
taxa was greatest in the Clifton region, then progressively less in the Toowoomba and
Dalby regions and least in the Cecil Plains region. This is proportional to the numbers of
localities in each region. There seems to be no obvious indication of regionalization
(except possibly for crocodilians as discussed below), so we tentatively suggest that the
eastern Darling Downs Pleistocene fauna is a single local fauna.
Rarity
The number of sites at which each taxon was recorded are given in Appendix 3. Of
the 68 taxa identified there, only 14 are found at more than five localities and only five
(Macropus titan, Protemnodon anak, P. roechus, ‘large Diprotodon’, ‘small diprotodon-
tid’) at more than 10. Thus there seem to be few widespread taxa and many restricted
ones, e.g., Palorchestes parvus, Sthenurus, Aepyprymnus, Propleopus, Onychogalea,
Osphranter and Thylogale which all seem to have been uncommon or restricted in range.
However, this may be misleading as large numbers of fossils have only vague locality
data such as “Darling Downs” so, for example, although only a single P. parvus is
recorded from any specific site, 11 other specimens (without specific locality data) are in
the QM collections. This example is the worst, and the other taxa here listed are repre-
sented by few or no (Aepyprymnus, Onychogalea, Osphranter and Thylogale) other
Pleistocene specimens from the Downs in the QM collections. Neither are they recorded
(at least from specific localities) by Lydekker (1887).
Proc. LINN. Soc. N.S.W., 117. 1997
126 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Although we regard those taxa found at several localities as having been wide-
spread animals on the Pleistocene Downs, “restricted” is used here to indicate taxa found
at few localities. We may interpret these taxa as having been uncommon, having had
small population sizes, and most are actually represented by few specimens in the collec-
tions. In fact, even the creatures with widespread distributions are represented by few
specimens, and the minimum number of individuals represented per locality is usually 1.
Whether the animals represented by uncommon fossils were rare in the sense of today’s
rare animals is a matter requiring taphonomic analysis.
Were these restricted taxa found all over the Downs in low numbers (actually rare) or
were they perhaps in large numbers restricted to specific locations, i.e., localized? If they
were widely distributed in low numbers we might expect to find them preferentially at
localities with large numbers of taxa (ranging to 25 taxa per locality), i.e., the larger sample
sizes. Of these restricted taxa (defined in this context as those found at 1—5 localities) some,
such as Macropus thor and Onychogalea unguifera are indeed found only in the large sam-
ple sizes. Only one, Phaps sp., is found only in a small sample size (<5) but others, such as
Procoptodon goliah and Megalania prisca, are found in both large and small sample sizes.
Being a bird Phaps, we think, was probably subject to both preservational and collecting
bias. Hence it may well have been more common than here indicated. So it seems likely
there weren't any localized populations but we would recommend more systematic collect-
ing, especially in the western regions, before drawing any further conclusions on this point.
Only a single human fossil, the Talgai skull, has been found on the Downs. Even
human artifacts contemporaneous with the megafauna are probably nonexistent as there
is only a single, unconfirmed report of their existence (Klaatsch, 1904), and we have
seen none in our collecting on the Downs. This may imply that the Downs fossils and
deposits date to a time before the entry of humans into Australia. Gill (1978) thought the
Talgai skull clearly postdated the extinct marsupials. Ninja turtles are also uncommon,
which is unexpected in view of their size and exuberant armor. In addition to the holo-
type of Ninjemys oweni, their presence is confirmed by a single meiolaniid vertebra from
near Macalister. This suggests that they were widespread — the holotype is from “Kings
Creek” — but quite rare.
Pliocene taxa
Several species considered to be Pliocene (e.g., by Archer, ef al., 1984) have been
recorded in the QM collections from the eastern Downs. These include Macropus dryas,
Osphranter woodsi, Protemnodon devisi, Euryzygoma dunense and possibly Macropus
pan. Four possibilities exist: 1) there are some unrecognized Pliocene deposits in the east-
ern Downs; ii) some Pliocene species persisted, perhaps in reduced numbers, into the
Pleistocene, ii1) they are simply mis-identified; and iv) the locality data are incorrect.
Even though detailed Pleistocene stratigraphy has yet to be carried out on the Downs, the
fact that all of putative Pliocene taxa derive from localities with large sample sizes
(17-25 taxa) suggests that the first possibility is unlikely. In order to very roughly assess
the likelihood of misidentification, we looked at the specimens of Euryzygoma reported.
These included an incomplete temporal from Macalister, nine isolated and worn incisors
(some of which may be macropod) from Freestone Ck. and Dalby and two dentaries with
very worn cheek teeth from Gowrie. Since we are not aware that diprotodont genera can
be accurately distinguished from fragmentary cranial elements or the roots of incisors we
feel that these identifications may be in error. From the amount of wear on the teeth —
which are usually considered the more diagnostic structures in mammals — in the
Gowrie specimens we hesitate to take these to be confidently identifiable. As for the
remainder of the taxa concerned, we cannot distinguish between the remaining possibili-
ties at this time, but our experience with the reported Euryzygoma material suggests that
misidentification cannot be ruled out as an explanation.
Proc. LINN. SOc. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ 127
Specific groups
Looking at the specific groups present on the eastern Downs, further comments are
warranted. Of the wombats, Phascolonus was the most widespread. This may be the result
of its large size, but the smaller Vombatus is also (moderately) widespread, suggesting that
both of these forms were actually prominent in the Downs mammalian fauna.
Potoroines and hypsiprymnodontines seem to have been rare but are also small, so
caution is advised. Of course if Propleopus was a predator, as has been suggested
(Archer and Flannery, 1985), its rarity is to be expected. Large macropodids, on the other
hand, were common; most sites have Macropus titan, Protemnodon anak and P. roechus.
Other macropods are found at fewer than ten sites, although M. agilis, M. ferragus and
Troposodon minor, found at eight to nine sites, are reasonably common. Sthenurines
(except 7: minor) are found at fewer sites, suggesting that they (even T. minor) were gen-
erally rarer than macropodines. But their presence does suggest that some of the eastern
Downs was wooded to a greater extent than when visited by Leichhardt in the early 19th
century.
Of the diprotodontids, the ‘large Diprotodon’ is found at almost all sites ranging
across the Downs up and into the dividing range. It is approximately eight times as com-
mon as the smaller form, which may imply that the smaller individuals were juveniles or
simply that the large and robust bones of larger animals were more easily preserved and
discovered than those of the smaller form. The smaller diprotodontids are still poorly
understood taxonomically so no conclusions about them are presently warranted.
Palorchestids seem only moderately widespread and monotremes seem to have
been rare. This is supported by our field experience — very few monotremes have been
found after much searching by both professionals and amateurs. As mentioned previous-
ly carnivores are expected to be rare. The most widespread were Thylacoleo and
Megalania, the latter confirming the significant role of reptilian predators in Australian
Pleistocene faunae.
Teleost fossils are rare and restricted to the eastern and southeastern regions. But in
our opinion, based on collecting experience, this is due to collecting bias. Teleost fossils
have been found at all of the systematically collected localities, except “Bundah’ (which
produced no certain fossils of aquatic or amphibious animals at all), and almost none of
the other sites. Dipnoans are a different matter, and have been known from the Pliocene
at Chinchilla since the nineteenth century. Those from the eastern Downs belong to the
same species, Ceratodus palmeri, as that from Chinchilla and are all recorded from
Kings Creek. We suspect that this accurately reflects their distribution when alive,
although why they have not been recorded from deposits on the Condamine, where they
lived during the Pliocene, is unknown. It may be that the Kings Creek population was a
relict population during the Pleistocene.
Absent taxa
Several forms that might reasonably have been expected are not in fact present in
the QM collections nor, to our knowledge, in any other collections from the eastern
Downs. Since these are animals of moderate to large size, we are reasonably confident
that they were actually absent, although we don’t know why. These are madtsoine snakes
(such as Wonambi), dromornithid birds and ziphodont crocodilians. The first two might
be expected on the basis of size — both emus and smaller snakes have been found —
and shed teeth of ziphodont crocs might be expected to have been collected, even if mis-
taken for teeth of Megalania. But none are present.
Possums do not occur in the QM (or London) collections from the Downs. This
might be attributed to their small size, except that they have not been found at systemati-
cally collected localities nor in searches specifically for them (Godthelp, pers. comm,
1995).
Proc. LINN. Soc. N.S.W., 117. 1997
128 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Crocodilians are more common and diverse in the Dalby region and this is borne
out by field experience. So while crocs were not actually absent in eastern regions they
seem to have been decidedly rare. Megalania, on the other hand, seems to have been
widely spread. It is not known from the Cecil Plains region, but this may be simply a
matter of the small sample from that region.
SUMMARY
There is presently no evidence for faunal regionalization or the existence of more
than a single local fauna in the eastern Darling Downs Pleistocene. Few taxa (14 out of
63) are found at more than five (of 68) localities and even fewer (five) at more than 10.
The uncommon taxa seem to have been widespread over the Downs, and may have had
small population sizes, or may have been subject to preservational bias. Sthenurines
seem to have been less common than macropodines, but their presence suggests that the
Pleistocene Downs did support some woodland. Ninja turtles almost certainly had small
populations. Species considered to be Pliocene are recorded, but whether they represent
remnant populations or simply mistakes is unknown. The large form of Diprotodon was
widespread and seems to have been common, whereas palorchestids were less common,
and monotremes seem to have been rare. The most common carnivorous forms were
Megalania and Thylacoleo. Crocodiles seem to have been most common and diverse in
the Dalby region. The rarity of teleost fossils is probably due to collecting bias, but lung-
fish seem to have been restricted to Kings Creek, perhaps as a relict population. There is
no indication that possums, ziphodont crocodiles, dromornithid birds or madtsoine
snakes inhabited the Pleistocene Downs.
ACKNOWLEDGMENTS
We appreciate the assistance of Bernard Cook, Angela Hatch, Henk Godthelp and Ian Sobbe in the
course of this research, and of Bevan and Debra Byers, John Mahon, Gary and Neville Rannfeldt, Julian
Ridgeway and Grantleigh Wolf in drawing our attention to and providing access to localities. Cornelia Kurz
was funded by the Deutscher Akademischer Austauschdienst.
REFERENCES
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Archer, M., Clayton, G. and Hand, S. (1984). A checklist of Australasian fossil mammals. In ‘Vertebrate
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Bartholomai, A. (1976). Notes on the fossiliferous Pleistocene fluviatile deposits of the eastern Darling Downs.
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Bennett, G. F. (1872). A trip to Queensland in search of fossils. Annals and Magazine of Natural History ser. 4,
9, 314-321.
Bennett, G. F. (1876). ‘Notes of rambles in search of fossils on the Darling Downs’. (Government Printer:
Australia).
French, M. (1989). ‘Conflict on the Condamine’. (Darling Downs Institute Press: Australia).
Gill, E. D. (1978). Geology of the late Pleistocene Talgai cranium from S.E. Queensland, Australia.
Archaeology and Physical Anthropology in Oceania13, 177-197.
Ingram, G. J. (1990). The works of Charles Walter de Vis, Alias “‘Devis’, alias ‘Thickthorn’. Memoirs of the
Queensland Museum 28, \—34.
Klaatsch, H. (1904). Ubersicht iiber den bischerigen Verlauf und die Errungenschaften seiner Reise in
Australien bis Ende September 1904. Zeitschrift fiir Ethnologie 1904, 211-213.
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Lydekker, R. (1887). ‘Catalogue of the fossil Mammalia in the British Museum (Natural History). Part V. The
Group Tillodontia, the Orders Sirenia, Cetacea, Edentata, Marsupialia, Monotremata and Supplement’.
(British Museum: England).
Lydekker, R. (1888). ‘Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natura? History).
Part I. The Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and
Protorosauria’. (British Museum: England).
Lydekker, R. (1889). ‘Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History).
Part III. The Order Chelonia’. (British Museum: England).
Macintosh, N. W. G. (1967). Fossil man in Australia. Australian Journal of Science 30, 86-98.
Mahoney, J. A. and Ride, W. D. L. (1975). Index to the genera and species of fossil Mammalia described from
Australia and New Guinea between 1838 and 1968 (including citations of type species and primary type
specimens). Western Australian Museum, Special Publication 6, 3-249.
Meltzer, D. J. and Mead, J. I. (1985). Dating late Pleistocene extinctions: theoretical issues, analytical bias, and
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U.S.A.).
Owen, R. (1845). ‘Descriptive and illustrated catalogue of the fossil organic remains of Mammalia and Aves
contained in the Museum of the Royal College of Surgeons of England’. (Richard and John E. Taylor:
England).
Owen, R. (1877). ‘Researches on the fossil remains of the extinct mammals of Australia; with a notice of the
extinct marsupials of England’. (J. Erxleben: England).
Van Tets, G. F. and Rich, P. V. (1990). An evaluation of de Vis’ fossil birds. Memoirs of the Queensland
Museum28, 165-168.
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Geology of Queensland’. Journal of the Geological Society of AustraliaT, 393-403.
Proc. LINN. Soc. N.S.W., 117. 1997
130 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
APPENDIX 1: FAUNAL LISTS FROM LYDEKKER (1887, 1888, 1889):
Clifton list includes ‘Clifton Plains’ and Kings Creek list, excludes those given as ‘King
Ck., Clifton’
Condamine River: Macropodinae
Diprotodontidae Macropus titan
Diprotodon, large form Osphranter altus
small diprotodontid
Thylacoleonidae
Thylacoleo carnifex
Crocodylidae
crocodilian
Varanidae
Megalania prisca
CECIL PLAINS REGION:
St. Ruth:
Diprotodontidae
Diprotodon, large form
Vombatidae
Phascolomys medius
Phascolonus gigas
Vombatus ursinus
Macropodinae
Macropus ferragus
Osphranter altus
Protemnodon brehus
Protemnodon roechus
Sthenurinae
Procoptodon rapha
Troposodon minor
TOOWOOMBA REGION:
‘Eton Vale’:
Vombatidae
Phascolomys magnus
Phascolonus gigas
Macropodinae
Osphranter altus
Protemnodon roechus
Diprotodontidae
Diprotodon, large form
Thylacoleonidae
Thylacoleo carnifex
Chelidae
chelid
Gowrie:
Vombatidae
Phascolomys magnus
Phascolomys medius
Phascolonus gigas
Vombatus ursinus
Proc. LINN. SOC. N.S.W., 117. 1997
Osphranter robustus
Protemnodon anak
Protemnodon roechus
Sthenurinae
Procoptodon rapha
Sthenurus atlas
Diprotodontidae
Diprotodon, large form
small diprotodontid
Dasyuridae
Dasyurus viverrinus
Sarcophilus laniarius
Thylacoleonidae
Thylacoleo carnifex
Crocodylidae
crocodilian
Varanidae
Megalania prisca
Hodgsons Creek:
Thylacoleonidae
Thylacoleo carnifex
Westbrook Creek:
Chelidae
chelid
CLIFTON REGION:
Clifton:
Vombatidae
Phascolonus gigas
Macropodinae
Macropus giganteus
Macropus titan
Protemnodon brehus
Protemnodon roechus
Sthenurinae
Procoptodon pusio
Diprotodontidae
Diprotodon, large form
Crocodylidae
crocodilian
Kings Creek:
Vombatidae
Phascolonus gigas
R.E. MOLNAR AND C. KURZ 1H
Macropodinae Varanidae
Protemnodon brehus Megalania prisca
Diprotodontidae Meiolantidae
Diprotodon, large form Ninjemys owenti
Thylacoleonidae
Thylacoleo carnifex
APPENDIX 2
Distribution of taxa on the eastern Darling Downs.
DR = Dalby region; CPR = Cecil Plains region; TR = Toowoomba region; CR = Clifton region.
DR CPR TR CR
Lasiorhinus latifrons/sp. X
Phascolomys angustidens Xx
P. medius X xX
Phascolonus gigas/sp. x x x
P. magnus X
Vombatus ursinus x X
V. sp. x
Aepyprymnus rufescens x
Propleopus oscillans x Ke
Macropus agilis X xX x
M. dryas x
M. ferragus X X X
M. giganteus X
M. pan (or faunus) x
M. pearsoni x x x x
M. piltonensis Xx
M. thor X x
M. titan X X x x
Onychogalea unguifera x x
Osphranter altus x x x
O. woodsi X
Procoptodon goliah X X
P. pusio xX x X
P. rapha X
Protemnodon anak x x X X
P. affinis X
P. brehus Xx X X
P. devisi x Xx
P. roechus x Xx X X
Proc. LINN. Soc. N.S.W., 117. 1997
12 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Sthenurus andersoni x x
S. pales
Thylogale sp.
Troposodon kenti
T. minor x x
Wallabia indra X Xx
Diprotodon small form x x x
Diprotodon large form X X XK
small diprotodontid X X X
Palorchestes azael xX x
P. parvus
Dasyurus spp.
Sarcophilus laniarius/sp. X x
Thylacinus cynocephalus x
Thylacoleo carnifex/sp. X X Xe
Perameles sp. e
Ornithorhynchus anatinus
rodent
Dromaius novaehollandiae x
Ardea cf. A. novaehollandiae
Ephippiorhynchus asiaticus
Phaps sp.
Progura gallinacea
Gallinula mortierii x
duck
quail
buteonine
Crocodylus porosus X
Pallimnarchus pollens xe
crocodilian X
Megalania prisca x xe
varanid
agamid ?
snake
meiolaniid Xx
chelid X x
trionychid
frog
Ceratodus palmeri
Tandanus tandanus
teleost x
Proc. LINN. SOc. N.S.W., 117. 1997
R.E. MOLNAR AND C. KURZ
APPENDIX 3
133
Ranking of taxa (mostly monospecific genera or species) by the number of sites at which they occur. Those
sites for which identifications were only to category higher than genus, or to genus for multispecific genera,
and “Jimbour Creek”, ““Condamine River” and “Kings Creek” were excluded, as being too extensive for single
localities in this context. 68 localities (not all with taxa identified to genus) are included. Numbers on the right
are the total number of taxa found at the sites at which the listed taxa occurred (given only for those that
occurred at fewer than 10 sites). *: species recorded as Pliocene in age.
Taxon No. of sites No. of taxa from each site
Lasiorhinus latifrons/sp. 2 17,1
Phascolomys angustidens 2 Zant
P. medius 2 22,1
Phascolonus gigas/magnus/sp. 8 25, 22, 18, 17, 14,4, 1,1
Vombatus ursinus/sp. 6 25, 19, 17, 16, 14,3
Aepyprymnus rufescens 1 25
Propleopus oscillans 1 9
Macropus agilis 8 25, 22, 19, 17, 14, 12,7,5
M. dryas * 1 25
M. ferragus 7 22, 19, 17, 16,9, 5,4
M. cf. M. giganteus 1 7
M. pan (or faunus) * 1 19
M. pearsoni 4 22, 16,7,5
M. piltonensis 1 19
M. thor 2 25, 19
M. titan 21 -
Onychogalea unguifera 2 D250
Osphranter altus 4 25, 17, 14,2
O. woodsi * 1 19
Procoptodon goliah 5 17, 14,9, 1,1
P. pusio 4 25,22, 17, 16
P. rapha 2 22, 14
Protemnodon affinis 1 17
P. anak 27 -
P. brehus 3 25, 22,3
P. devisi * 2-3 25, 19?, 17
P. roechus 12 -
Sthenurus andersoni 4 25, 22, 17,3
S. pales 1? 19
Thylogale sp. 1 22
Troposodon kenti 1 12
T. minor 8 25, 22, 19, 17, 16, 12,7, 1
Wallabia indra 2 25, 12
Diprotodon small 5 17, 16,5, 4, 1
Diprotodon large 39 =
small diprotodontid 15 —
Proc. LINN. Soc. N.S.W., 117. 1997
134 DISTRIBUTION OF PLEISTOCENE VERTEBRATES
Palorchestes azael
P. parvus
P. sp.
Dasyurus sp.
Sarcophilus laniarius/sp.
Thylacinus cynocephalus
Thylacoleo carnifex
Perameles sp./bandic.
Ornithorhynchus anatinus
rodent
Dromaius novaehollandiae/sp.
Ardea cf. A. novaehollandiae
Ephippiorhynchus asiaticus
Phaps sp.
Progura gallinacea
quail
Gallinula mortierii/sp.
Crocodylus porosus
Pallimnarchus pollens
crocodilian
Megalania prisca
varanid
agamid
snake
meiolaniid
chelid
trionychid
frog?
Tandanus tandanus
teleost
Proc. LINN. SOC. N.S.W., 117. 1997
MN Ss OM OW = w
SJ Ay Ss OS OR OO eS) ULES OOO OS OO eS Ee SS SS Lh ULE YDS) h®D
22, 16, 12, 5,2
1
18,1
22, 18, 17, 14, 12
17,9,9,9
25,22, 17,9
25, 17, 17, 12,9,9,7,5,4
18, 14,6
17
18,9
17, 12
14
19
25, 19, 18, 12, 12,7
22, 19, 18, 17, 12, 12,4, 1
18, 12
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18
16
25, 18, 16, 12, 12,7,3
19
18
22
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Analysis of a Late Quaternary Deposit and
Small Mammal Fauna from Nettle Cave,
Jenolan, New South Wales.
DEBORAH A. Morris'!, M.L. AUGEE', D. GILLIESON? AND J. HEAD’.
‘School of Biological Science, University of NSW, Sydney NSW 2052;
*School of Geography and Oceanography, University College, University of NSW,
Australian Defence Force Academy, Canberra ACT 2600;
‘Radiocarbon Laboratory, Australian National University, Canberra ACT 2601.
Morris, D.A., Augee, M.L., Gillieson, D. and Head, J. (1997). Analysis of a late Quaternary
deposit and small mammal fauna from Nettle Cave, Jenolan, New South Wales.
Proceedings of the Linnean Society of New South Wales 117: 135-162
A deposit of small mammal bones in Nettle Cave, part of the Jenolan Caves system,
was excavated. The bone deposit appears to be the result of owl pellet accumulation. A pair
of Sooty Owls (Tyto tenebricosa) currently inhabits a roosting site within the cave. The
deposit was excavated to a depth of 68 cm, which represents an accumulation throughout the
last glacial recession in the late Pleistocene to the present. Two radiocarbon dates (7,140 +
280 and 8,730 + 280 BP) were obtained from discrete charcoal lenses in the middle layers of
the deposit. Analyses of small mammal remains and sediments indicate climatic conditions
during the late Pleistocene were colder and drier than at present, becoming warmer and wetter
in the Holocene. The apparent abrupt extinction of Burramys parvus and the rapid decline in
abundance of Mastacomys fuscus in the Jenolan area are attributed to a brief humid period
that occurred in southeastern Australia at around 15,000 to 14,000 BP.
Manuscript received 1 June 1996, accepted for publication 23 October 1996.
KEY WORDS: Nettle Cave, owl pellet accumulation, Tyto tenebricosa, late Quaternary,
Burramys parvus, sediments, climate.
INTRODUCTION
Fossil deposits have long been used as a basis for the interpretation of past envi-
ronments. In Australia, various Quaternary cave deposits have been analysed and used to
reconstruct the faunal and climatic history of the surrounding area (e.g., Balme et al.
1978, Baynes 1987, Baynes et al. 1976, Hope et al. 1977, Porter 1979, Wakefield 1972).
Deposits containing small mammal remains have been found to be particularly useful
(Lundelius 1963).
Little fossil material had been found in Jenolan Caves until a collection of bones was
analysed from a small cave overlooking the lower carpark (Hope 1979). Infrequent discov-
eries of isolated skeletal remains from various caves have been reported in subsequent
years. This study involved excavation of a fossil deposit in Nettle Cave, part of the Jenolan
Caves system. The excavation yielded an abundance of intact small mammal bones, includ-
ing those belonging to now locally extinct taxa, together with avian postcranial material
and a few agamid and scincid mandibles (which are not discussed in this paper).
MATERIALS, METHODS AND STUDY AREA
The Jenolan Caves Reserve is situated on the Great Dividing Range (33°47’S,
Proc. LINN. Soc. N.S.W., 117. 1997
136 LATE QUATERNARY DEPOSIT
entrance
Figure 1. Plan of Nettle Cave, Jenolan, with the area of heaviest deposition of owl pellets outlined (lower cave).
The highlighted area is the quadrat selected randomly for excavation. This area was further subdivided into
quarters as shown in the insert. The quadrat and the subdivided areas are not to scale. The current Sooty Owl
roost is indicated by the arrow (upper cave). (modified from Cox et al. 1989).
150°02’E; 1,100—1,200 m a.s.l.) approximately 110 km west of Sydney. This area has a
maximum yearly mean temperature of 16.6°C and a minimum of 8.0°C (recorded at
Katoomba - approximately 30 km from Jenolan). The average annual precipitation is
1,412 mm with the greatest rainfall occurring between December and June. The vegeta-
tion within the Jenolan Caves Reserve supports eight major vegetation communities
(Lembit 1988) ranging from open forest to cleared land.
Nettle Cave is a high-level entrance into the Devil’s Coach House, which 1s itself a
natural tunnel approximately 80 m high and 40 m wide (Cox et al. 1989). Fiowstone
forms a false floor in Nettle Cave (Anon. 1988). Cave conditions are dry (Nettle Cave is
about 20 m above modern flood levels), with an annual temperature range from below
zero to 30°C (Cox et al. 1989). The cave receives light from the entrance in the south,
from a roof-hole in the northeast and from Arch Cave in the southwest (Fig. 1).
The fossil deposit examined in this study is concentrated beneath a rock ledge in
the roof of Nettle Cave close to the northern wall of the lower cave (Fig. 1). A pair of
Sooty Owls (Tyto tenebricosa) currently occupies a nocturnal roosting site in the north-
facing wall of the upper Nettle Cave (Fig. 1). Sooty Owls were first reported roosting in
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 137
Loose powdery soil
—
& Bone-rich sediments
1)
—
3
& 10
fmt
=}
Nn Dark organic layer with
[-) carbonate cement
va
S Pie
Oo fas) Brown gritty sandy clay
(0.05).
TABLE | Radiocarbon ages of charcoal from the Nettle Cave deposit.
Sample Depth dl4C 613C D14C age
code (cm) (permil) (permil) (permil) (years BP)
ANU-7897 28-29 - 587.8 + 13.7 - 24.0 + 2.0 - 588.6 + 13.8 7,140 + 280
ANU-7898 35-36 - 661.9 + 11.4 - 24.0 + 2.0 - 662.6 + 11.5 8,730 + 280
Source of the Sediment
Mineral magnetic results from the Nettle Cave sediment samples are compared
with data from other sites in the Jenolan Caves catchment (Stanton 1989) in Table 2. The
results indicate that the Nettle Cave sediments are distinct from these possible sources
and especially from the Mammoth Cave fluvial sediments.
TABLE 2 Mean magnetic parameters for different sediment source areas, Jenolan catchment. SIRM denotes
saturation isothermal remanent magnetisation. Data other than that for Nettle Cave from Stanton (1989).
Mean Mean frequency Mean
Source area susceptibility dependent susceptibility SIRM
Nettle Cave 1.43 9.9 20.6
Terrace Creek 4.24 6.9 41.4
Jenolan-Bindo 4.4] VS 49.6
Creek Divide
Western Ridge 9.24 4.2 109
Mammoth Cave 4.24 5.4 39.9
Fines
Mammoth Cave Syoil ae B25)
Coarse
Particle size analysis can provide some information on sediment transport and
deposition (Krumbein and Sloss 1963). The relative amounts (by weight) of gravel, sand,
silt and clay in each sample from the Nettle Cave deposit are illustrated in Fig. 3.
(Unfortunately, bone was included in the gravel fraction; if bone had been excluded, the
relative proportions of sand, silt and clay would be greater than that suggested in Fig. 3.)
The greatest fluctuations, albeit minor, were in the proportions of gravel and then sand.
An increase in the amount of fine material occurred around 37 cm. This increase was
coincident with water ponding of the top layer in the lower zone of the deposit. The pre-
ponderance of coarser, angular particles in the sediment samples suggests that the sedi-
ment is of local origin and has not been subjected to lengthy fluvial transport.
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 141
SEDIMENTS FROM THE DEPOSIT (%)
DEPTH INTERVAL (CM)
Figure 3. Graphical representation of the amount of gravel, sand, silt and clay (by weight) in the sediments
from the Nettle Cave deposit.
Considering the mineral magnetic data and particle size of the Nettle Cave sediments,
these sediments are likely to have originated from local soil above the cave. Moreover,
the relatively minor fluctuations in the proportions of the sediments throughout the depth
of the deposit suggest there was little variation in the source of the sediments.
Mammal Fauna from the Nettle Cave Deposit
Thirty-five species of mammals were identified in the Nettle Cave deposit
(Appendix B). Of these species, 74% are extant and 26% are extinct either locally or in
southeastern Australia.
If a species is present in an upper or lower level of the deposit, it tends to appear
throughout the upper or lower zone of the deposit, respectively (Table 3). The distribu-
tion of some species at 41 to 44 cm appears to be discontinuous.
The majority of the species occurring throughout the deposit (Table 3) are small
animals weighing less than 200 g. All these specimens are adult. The larger species pre-
sent, e.g. Dasyurus sp., P. peregrinus, Isoodon obesulus and Perameles nasuta, are rep-
resented by subadullts.
Species with restricted habitat requirements and range of distribution are the most
useful indicators of environmental conditions (Baynes et al. 1976). The changes in rela-
tive abundance with time of selected non-volant, small mammal species from the Nettle
Cave deposit are illustrated in Fig. 4. These species were selected as indicators of possi-
ble environmental change in the Jenolan area for the following reasons:
1. the species showed a change in distribution over time or,
2. the species showed a change in abundance over time and
Proc. LINN. SOc. N.S.W., 117. 1997
142
LATE QUATERNARY DEPOSIT
TABLE 3 Summary of Appendix B: Presence or absence of specimens identified from each depth interval from the
Nettle Cave deposit. Presence of species in a depth interval is indicated by a black block. Depth intervals 13-5 cm and
41-39 cm were not analysed, and are represented as narrow, blank bars. S = number of non-volant mammal species in
the deposit. n=sum of the MNIs of non-volant mammal species, excepting A. spp., S. sp., P. spp. and R. spp. 1.
Presence of bats in a depth interval is indicated by a dashed block, but the MNI is not included in n.
2. Presence of
birds is indicated by a dashed block; specimens were not identified above order. 3. Skinks were present in depth
intervals 5-0 cm and 25—13 cm; agamids were present in depth intervals 5—O, 25-13, 35-25, 37-35 and 39-37 cm.
SPECIES
Antechinus stuartii sensu lato
A. swainsonit
A. flavipes
A. spp.
Sminthopsis murina
S. sp.
Phascogale tapoatafa
Dasyurus sp. cf. D. viverrinus
Isoodon obesulus
Perameles nasuta
Pseudocheirus peregrinus
Petauroides volans
Petaurus breviceps
Cercartetus nanus
C. lepidus
Burramys parvus
Acrobates pygmaeus
Potorous sp. cf. P. tridactylus
Bettongia sp.
Thylogale thetis
Conilurus albipes
Pseudomys oralis
P. gracilicaudatus
P. australis
P. novaehollandiae
P. fumeus
P. spp.
Mastacomys fuscus
Rattus fuscipes
R. rattus
R. spp.
Mus musculus
Oryctolagus cuniculus
Bats
Birds?
Lizards
Proc. LINN. SOC. N.S.W., 117. 1997
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3. the species’ habitat requirements are both well-documented and restricted.
Modern Mammal Fauna in the Jenolan Area
The term ‘modern mammal fauna’ is used to describe the fauna inhabiting, or
thought to inhabit, the Jenolan area since European settlement. Of the 35 mammal
species recovered from the Nettle Cave deposit (Table 3), one (Conilurus albipes) is pre-
sumed to be extinct (Watts and Aslin 1981); eight (Dasyurus viverrinus, Cercartetus lep-
idus, Burramys parvus, Bettongia sp., Pseudomys fumeus, Pseudomys australis,
Pseudomys oralis and Mastacomys fuscus are extinct in the area (Strahan 1995; Watts
and Aslin 1981); Pseudomys gracilicaudatus had previously occurred this far south
(remains were found in superficial deposits at Walli Caves near Canowindra and
Wombeyan Caves [Mahoney and Posamentier 1975]); the status of Phascogale tapoatafa
and Pseudomys novaehollandiae in this area is uncertain (Strahan 1995; Watts and Aslin
1981); the remaining 23 are locally extant (Strahan 1995).
DISCUSSION
Age of the Deposit
The age of specimens in the Nettle Cave deposit can be estimated when associated
with stratigraphy, other fauna, radiometric dates and the appearance of the material (after
the manner of Baynes (1987)).
Two radiocarbon dates based on charcoal samples were obtained from the middle
levels of the deposit (Fig. 2). If one assumes a constant rate of accumulation of sediments
to the base of the excavation at 68 cm, then this level may represent 16,000—14,000 years
BP. However, ponding and a change in the nature of sedimentation is evident below
35-36 cm (Fig. 2). In addition, abrupt changes in the faunal assemblage around 41-43
cm may indicate either a hiatus in deposition, or a minor unconformity (period of nonde-
position or erosion) in the deposit (Krumbein and Sloss 1963). Conversely, changes in
the composition of the material being deposited may have produced the change in strati-
fication (Dunbar and Rodgers 1963). For example, changes in grain size may cause pro-
nounced layering. Therefore, this time frame on the basis of sedimentation should be
treated with caution, since charcoal was not available at the base of the pit to allow more
precise dating.
Environmental History at Jenolan based on Nettle Cave Sediments
Although real precipitation at the end of the Pleistocene was reputedly low (Dodson
1977; Galloway 1965), seasonal melting of the snow would have made available free
water. A study by one of us (D.G., unpublished data) suggests the influx of subsoil parti-
cles to the lower levels of the Nettle Cave deposit indicates erosion of the topsoil overly-
ing the cave either due to hillslope instability (Gillieson et al. 1985) or thawing of the
ground and subsequent washing away of this surface material. Wind activity would have
contributed to the erosion to some degree. More importantly, wind activity is a selective
barrier in the transport of particular grains. Coarse particles such as gravel are left behind
or deposited close to the source forming a local accumulation. Fine grains (silt and clay)
are kept in suspension and transported over long distances (Krumbein and Sloss 1963;
Pettijohn 1957; Reineck and Singh 1975). These conditions are reflected in the relatively
low amounts of silt and clay in the Nettle Cave sediments (Fig. 3).
The sediment in the levels of the deposit around 44-41 cm consists of a yellow-
brown, gravelly clay which was probably the result of local water ponding in the cave.
Proc. LINN. Soc. N.S.W., 117. 1997
144 LATE QUATERNARY DEPOSIT
A)
50 +
45 +
40 +
BA. pygmaeus
35 =
OP. volans
SSP. breviceps
BP. peregrinus
RELATIVE ABUNDANCE (%)
N
nn
04 f i, E. |
L12 L10 L8
L16 L14
DEPTH INTERVAL
B)
a
oO
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>
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OA. swainsonii
Ww
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BC. nanus
BSB. parvus
EAC. lepidus
w
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+
20 +
40 -
5 +
0 | EA ss | L = ac il 0 peeeen $— ae | jhe Oe Oke + _, Oe ++ i
L16 L14 L12 L10 L4
L8 L6 L2 SURFACE
DEPTH INTERVAL
RELATIVE ABUNDANCE (%)
nN
a
+
Figure 4. The relative abundance of selected non-volant, native mammal species from the Nettle Cave deposit.
The species represented are possible indicators of climatic change. Levels L2 and L7 were not analysed. See
Appendix B for the corresponding depth interval to each level. Continued on following page.
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 145
Cc)
50 +
45 + SSE Sree ees
|
| OP. oralis |
40 + | |
| SM. fuscus |
| |
| }
_ 35+ N | BR. fuscipes |
g . 3
z 30 + \ N \ .
S * fF FS \
z N \ N N : N
L14 L12 L10 L8 L6 L4 L2 SURFACE
DEPTH INTERVAL
Figure 4. Continued from previous page.
Ponding may have occurred from surface water seeping into the limestone and accumu-
lating behind the cave wall. Numerous runnels on the wall behind the excavation suggest
water inflowing. Leaching to the lower layers would have followed. An increase in the
amount of fine material (Fig. 3) at these levels corresponds with more humid conditions,
a phenomenon reported by Wells et al. (1984) for Victoria Cave. Consolidation of parts
of these levels may have been due to the introduction of cementing material in the early
stages of diagenesis, rather than from compaction (Krumbein and Sloss 1963). In fact,
alkaline conditions and an increase in temperature would enhance the precipitation of
calcium carbonate which in turn would promote cementation of the sediment (Pettijohn
1957; Reineck and Singh 1975).
The radiocarbon dates of 7,140 + 280 and 8,730 + 280 BP agree well with the sedi-
mentary analyses. Between the two charcoal lenses was a white, calcareous, cemented
layer (Fig. 2). Its formation appears to have coincided with the wettest period at 7,500 to
5,000 BP (Bowler et al. 1976). Relative increases in the proportion of fine particles at 37
cm (Fig. 3) indicate wetter conditions. These conditions seem to correlate with the early
to mid-Holocene humid period (Bowler et al. 1976; Colhoun et al. 1982). Drier condi-
tions throughout the remainder of the Holocene (Bowler et al. 1976; Hooley et al. 1980)
are reflected in the proportionately low amounts of clay and silt in the upper levels of the
Nettle Cave sediments (Fig. 3). However, particle size analysis of these sediments indi-
cates the drier conditions of the late Holocene were still wetter than the conditions expe-
rienced in the late Pleistocene.
Mode of Deposition
Sooty Owls are the current source of small mammal bone deposits in Nettle Cave.
We consider that owls of some species have been the source of the deposit over the entire
Proc. LINN. Soc. N.S.W., 117. 1997
146 LATE QUATERNARY DEPOSIT
period of deposition covered by our excavation for the following reasons:
1. Long bones are intact; skulls are either intact (except for a damaged or absent
occiput) or broken into their component bones; mandibles are rarely heavily fragmented;
and little erosion of bone and tooth has taken place.
2. There is a bias towards smaller animals, particularly murids and small dasyurids;
the larger species are represented by subadults or juveniles.
3. Evidence for collection by diurnal raptors, e.g. kestrels, such as heavy digestive
erosion of bones and tooth enamel (Andrews 1990; Kusmer 1990) is absent. Evidence
for mammalian carnivores, e.g. Sarcophilus or Thylacinus, such as highly fragmented
bones (especially skulls), eroded teeth and bone-bearing coprolites (Lundelius 1966) is
also absent.
4. There is no evidence for fluvial deposition from the sedimentological data.
While the material from the Nettle Cave deposit fulfils the requirements of an owl
accumulation (Lundelius 1966), it is impossible to conclude that Sooty Owls have been
responsible for the entire deposit. Occupation of the roosting site may have alternated
between Masked Owls (Tyto novaehollandiae) and Sooty Owls for the duration of pellet
deposition. Both species have been found living in caves (Anon. 1988; J. Calaby, pers.
comm., in Wakefield 1960; pers. obs.) and take similar prey items of comparable sizes
(Table 4). Powerful Owls (Ninox strenua) may also have contributed to the deposit.
However, the Powerful Owl habitually roosts in trees rather than caves (Fleay 1944) and
is a more specialist predator, taking mid- to large-size arboreal species almost exclusive-
ly (Fleay 1944; James 1980; Kavanagh 1988; Tilley 1982) (Table 4). The almost total
absence of moderate-size arboreal mammals in the lower levels of the deposit (Table 3),
prey known to be taken by Sooty Owls and Masked Owls, suggests a lack of trees in the
Jenolan area at this time. A lack of trees would also make it unlikely that Powerful Owls
would have been present, given their habitat preference for forests (Kavanagh 1988) and
therefore, they were unlikely to have contributed to the Nettle Cave deposit.
Environmental History at Jenolan based on Mammal Remains
The Nettle Cave faunal assemblage containing B. parvus (Table 3) appears to be
late Pleistocene in age. The presence of B. parvus together with M. fuscus and
Antechinus swainsonii throughout this early phase of the deposit indicates a colder envi-
ronment than at present in the Jenolan area (Table 5). Modern B. parvus is physiologi-
cally intolerant of high temperatures (Fleming 1985) and is restricted to high alpine
areas in NSW and Victoria. The presence of these species and the virtual absence of
arboreal mammals suggest the vegetational formation was dominated by open areas
with a dense ground cover of grasses and low shrubs. Conversely, the low representa-
tion of arboreal mammals may be due to a smaller owl as predator, rather than a lack of
trees. However, the absence of A. pygmaeus and the low abundance of Petaurus brevi-
ceps in the lower zone of the deposit probably reflects the available vegetation at the
time rather than predator bias, given that owls will take arboreal mammals if they are
present (Table 4). The presence of M. fuscus may indicate the immediate presence of
water (Hope et al. 1977), either as snow, running streams or wet microhabitats in the
grasslands. Small pockets of forest or woodland were likely to have been pyesent at
lower altitudes.
The faunal composition of the lower levels of the Nettle Cave deposit appears to
broadly correspond to the assemblage described in the Pleistocene fraction of the
Pyramids deposit at Buchan (Wakefield 1969, 1972) and the entire Wombeyan breccia
(Ride 1960). The relatively higher abundance of M. fuscus and B. parvus and lower
abundance of A. swainsonii and arboreal species in the lower Nettle Cave deposit com-
pared with the relative abundances of these species in the older fraction of the Pyramids
deposit, suggest a less heavily forested vegetation in the Jenolan area, although sampling
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 147
TABLE 4 A list of the recorded prey items of the Sooty Owl, Masked Owl and Powerful Owl from the litera-
ture. a = adult, j = juvenile, yes denotes that the species is taken as prey by that particular owl, ? = this prey is
possibly taken. 1. Howe (1935); Hyem (1979); Loyn et al. (1986); Schodde and Mason (1980); Smith (1984). 2.
Hyem (1979); Mooney (pers. comm.). 3. Hyem (1979); James (1980); Kavanagh (1988); Seebeck (1976); Tilley
(1982).
Sooty Owl! Masked Owl’ Powerful Owl’
Prey species (Tyto tenebricosa) (T. novaehollandiae) (Ninox strenua)
Petaurus breviceps yes yes (a, j) yes (a)
P. australis yes
Pseudocheirus peregrinus yes (a, j) yes (a) yes (a, j)
Petauroides volans yes (j) yes
Trichosurus vulpecula yes (J) yes (j)
Cercartetus nanus yes yes (a)
C. lepidus yes (a)
Acrobates pygmaeus yes
Antechinus stuartii yes yes
A. swainsonii yes yes
A. minimus yes (a)
Sminthopsis leucopus yes
Phascogale tapoatafa yes
Dasyurus maculatus yes (a)
D. viverrinus yes (a, j)
Isoodon obesulus yes (j)
Perameles gunnii yes (a, j)
Bettongia gaimardi yes (a)
Potorous tridactylus yes (a)
Thylogale thetis ?
T. billardierii yes (j)
Hydromys chrysogaster yes (a)
Mastacomys fuscus yes yes (a)
Pseudomys fumeus yes
P. higginsi yes (a, j)
Rattus fuscipes yes yes (a)
R. lutreolus yes (a)
R. rattus yes yes (a)
R. norvegicus yes (a)
Mus musculus yes (a)
Felis cattus yes (j)
Lepus capensis yes (j)
Oryctolagus cuniculus yes () yes
Bats yes yes
Insects 2 yes yes
Birds yes yes yes
Frogs yes
Proc. LINN. Soc. N.S.W., 117. 1997
LATE QUATERNARY DEPOSIT
148
87 81 é sok sok sok sok
87 97 ‘FI “9 00S*Z-00S'I sah sak sak
87 ‘97 ‘*L ~=—006' 1-006 sok sok sok sak
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1997
Proc. LINN. SOC. N.S.W., 117.
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 149
Mt Kosciusko
2000 Barrington Tops
G
ir)
5
3 1500
2 1000
& TIMBER LINE
3S
=A 500 = Wombeyan
‘Oo Caves
0
32 33 34 35 36 37 38
Latitude (°S)
Figure 5. An altitude profile from Barrington Tops to Bass Strait with the location of Burramys parvus fossils
from the Pleistocene, and the estimated position of the tree-line and permanent snowline during the height of
glaciation at that time. However, the position of the snowline should be higher. The positions of the Buchan and
Wombeyan Caves have been lowered from that indicated by Caughley (1986) to that stated by Wakefield (1969).
The scale on the ordinate axis has been lowered so Bass Strait is at sea level. (modified after Caughley 1986).
bias from different predators at the two sites cannot be ruled out. The higher altitude of
Nettle Cave lends support to the former scenario. Caughley (1986) estimated the position
of the tree-line at the height of glaciation during the late Pleistocene to be below the
Jenolan, Wombeyan and Buchan Caves (Fig. 5). However, altitudinal data from
Wakefield (1969) indicate the tree-line would have been above the Buchan and
Wombeyan Caves at this time. While periglacial activity did not reach the southern
flanks of the Blue Mountains (Peterson 1968), the ground may have been seasonally
frozen (Gillieson et al. 1985). The apparent presence of trees at Jenolan indicates the
tree-line had shifted up and climatic warming in the highlands of southeastern Australia
had begun. Wakefield (1969, 1972) identified the older fraction of the Pyramids deposit
as having accumulated over the period covering most of the glacial recession phase at the
termination of the Pleistocene. This agrees well with the climatic and vegetational
sequences indicated by the faunal composition and sedimentological data of the lower
Nettle Cave deposit.
The age of the Nettle Cave deposit cannot be estimated with any certainty on the
rate of sedimentation alone. However, from the faunal evidence and comparison with the
Pyramids deposit and the climatological data from southeastern Australia (Bowler et al.
1976; Galloway and Kemp 1981), the lower levels of the Nettle Cave deposit appear to
have accumulated after the glacial maximum, about 20,000 years ago (Frakes et al.
1987), to around 15,000 BP.
Following this early phase was a transition period, represented by approximately
44-41 cm of the deposit, and correlating with the period 15,000 to 10,000 BP. This peri-
od was marked by an abrupt change in the composition and/or abundance of the faunal
assemblage in the deposit (Table 3, Appendix B) along with a change in the sediments
(Fig. 3). Burramys parvus and C. lepidus were no longer represented, the abundance of
A. swainsonii and M. fuscus was greatly reduced while the numbers of Rattus fuscipes, P.
peregrinus, P. volans and P. breviceps suddenly increased. While absence from the
deposit and discontinuity of distribution of some species at around 44—41 cm (Table 3)
may indicate a change of predator, the presence of other species throughout the deposit,
such as Dasyurus sp., I. obesulus, P. nasuta and most native murids, suggests changes in
Proc. LINN. Soc. N.S.W., 117. 1997
150 LATE QUATERNARY DEPOSIT
abundance of most species were the result of changing climatic conditions.
As previously mentioned, B. parvus is physiologically intolerant of high tempera-
tures (Fleming 1985). The presumed abrupt local extinction of B. parvus may therefore
indicate a sudden increase in temperature. The zoogeographic range of B. parvus may
have started to narrow during the brief period of humidity at 15,000 to 14,000 BP
(Bowler et al. 1976). The presence of A. swainsonii and M. fuscus indicates cold condi-
tions (Table 5). The lower abundance of these species in the 44-41 cm of the deposit
suggests warmer conditions than previously experienced. An increase in abundance of R.
fuscipes occurred at around the same time (Appendix B). Changes in vegetation at the
close of the Pleistocene from grasslands, with grasses being a major food source for M.
fuscus, to a ground cover dominated by ferns and shrubs, the preferred habitat of R.
fuscipes, may have given R. fuscipes a competitive advantage, thereby maintaining a
lower population of M. fuscus in the Jenolan area throughout the Holocene.
The local extinction of C. lepidus cannot be explained by climatic change alone.
The modern distribution of this species includes hot and dry conditions (Aitken 1977;
Dixon 1978). The coexistence of B. parvus and C. lepidus throughout the late
Pleistocene suggests the latter species is capable of tolerating climatic extremes. The dis-
appearance of C. /epidus from the Jenolan area may be attributed to a possible lack of
floristic diversity at the time. It has been claimed that modern C. lepidus is highly
mobile, and follows plants as they flower throughout the year (Ward 1992). As the
Jenolan area became more heavily forested towards the end of the Pleistocene, C. lepidus
may have been replaced by its larger congener, C. nanus.
The presence of arboreal species in the upper levels of the deposit suggests the
presence of open forest and woodland. These vegetational communities were well
established by 11,500 to 9,000 BP (Kershaw 1981). A combination of increased real
precipitation (Bowler et al. 1976; Kershaw 1981) and an increase in temperature
(Binder and Kershaw 1978) around this period may have contributed to the forestation.
Increased wetness is evident from the nature of the sediments in the deposit (Fig. 3),
while the continued low representation of A. swainsonii and M. fuscus is suggestive of
warmer conditions.
The upper levels of the Nettle Cave deposit (c. 41— 0 cm) appear to have accumu-
lated throughout the Holocene. The mammal species in the deposit indicate both wet and
dry sclerophyll forest and woodland with varying amounts of ground cover (Table 5).
The vegetation in the Jenolan area throughout the Holocene therefore appears to have
been much the same as that found in the area today.
The upper levels of the Nettle Cave deposit broadly resemble the younger fraction
of the Pyramids deposit. The species composition from this fraction of the Nettle Cave
deposit gives no indication of an arid period, as proposed by Wakefield (1969, 1972) for
the Pyramids deposit, but rather a change to perhaps slightly drier conditions. The faunal
assemblage from the final phase of the transition period in the Nettle Cave deposit is
comparable with the assemblage from the most recent Pleistocene sediments of the
Pyramids deposit.
The cave surface mammal fauna (Table 3) closely resembles the modern mammal
fauna in the Jenolan area with a few notable exceptions. Introduced species are found
with species no longer extant in the area (P. oralis and M. fuscus) or extinct (C. albipes).
Mixing of the deposit, possibly by rock-wallabies, or humans when the cave was open to
tourists, may have occurred at the surface and top level of excavation (0-5 cm). Mixing
is quite likely because the appearance of the bone from the surface differs from white to
pale grey for the introduced species, to a yellow-cream for the native species. The colour
of the bones from 0-5 cm varies from pale to dark grey for the introduced species, com-
pared with yellow-cream to dark orange for the native murids. Wakefield (1972) con-
cluded that variation in colour is correlated with age, whereby the lighter coloured bones
represent a more recent age than the darker coloured bones. Differential colouration of
Proc. LINN. SOc. N.S.W., 117. 1997
151
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD
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Proc. LINN. Soc. N.S.W., 117. 1997
152 LATE QUATERNARY DEPOSIT
bone occurred according to the depths at which the bones were initially deposited. The
presence of different coloured bones together suggests mechanical mixing of layers after
colour alteration of the bones had taken place, that is, the darker, older bones from a
lower level were mixed with the lighter coloured, younger bones from a more superficial
level. Therefore, by using Wakefield’s (1972) colour-age criterion, it may be assumed
that there was limited temporal overlap between these introduced and native species.
However, this assumption should be treated cautiously given that stratigraphic control of
the excavation was not sufficient to detect this overlap. Therefore, C. albipes, P. oralis
and M. fuscus may have been contemporaries of the introduced species. It is possible that
the local extinction of these three native mammals was a result of the introduction of
non-native mammals associated with European settlhement (Wakefield 1960; Watts and
Aslin 1981), e.g., rodents and rabbits. Distributional data on P. australis (Watts and
Aslin 1981), P. novaehollandiae (Keith and Calaby 1968; Strahan 1995) and P. fumeus
(Watts and Aslin 1981) in conjunction with the colour of the bones of these species from
0-5 cm, suggest they were not contemporaneous with the introduced species.
On the basis of the Nettle Cave data, major extinctions of the small mammal fauna
in the Jenolan area at the end of the Pleistocene do not appear to have occurred. Two
species (B. parvus and C. lepidus) probably became locally extinct at this ttme. The most
parsimonious explanation for most of the faunal changes at the end of the Pleistocene is
that of climatic change, particularly an increase in temperature. Local extinction of M.
fuscus, P. oralis and C. albipes at the time of European settlement was possibly the result
of competition from introduced species.
A Summary of the Chronology of Events
The available geomorphological and palaeontological evidence from the Nettle
Cave deposit and the proposed correlation with other events is summarised in Table 6.
The small mammal assemblage represented in the lower Nettle Cave deposit (68—
c. 44 cm) appears to be late Pleistocene in age. We suggest this fraction of the deposit
was accumulated during the glacial recession at the terminal phase of the Pleistocene, i.e.
after 20,000 to 15,000 BP. Conditions were colder and with less real precipitation than at
present. The vegetation was dominated by shrubs and a dense ground cover, with pockets
of wet sclerophyll forest. The disappearance of B. parvus from the Nettle Cave deposit is
attributed to the increase in temperature in southeastern Australia over the period 15,000
to 14,000 BP. A brief hiatus in deposition may have followed. The appearance of arbore-
al species may signify the revegetation of the area by forest and/or woodland. This event
corresponds with the period of increased precipitation from 11,500 to 9,000 BP. This
transition period of increased wetness is represented by ponding at roughly 44-41 cm of
the deposit.
The small mammal assemblage represented in the upper levels (41-0 cm) of the
Nettle Cave deposit is identified as Holocene. A wet forest fauna is suggested to have
inhabited the area during the early to mid-Holocene. Drier conditions, much the same as
the modern vegetational communities in the Jenolan area today, followed this early
Holocene humid phase. However, this period was wetter than that in the late Pleistocene.
The surface of the deposit is modern. Mixing with the upper levels of the Holocene
fraction occurred. Modern specimens could be distinguished from specimens deposited
prior to European settlement largely on the basis of discolouration of the bone. Local
extinctions, most notably of rodents, possibly occurred after European settlement as a
consequence of competition from introduced mammals.
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 153
ACKNOWLEDGEMENTS
Our thanks go to Dr Suzanne Hand for identification of the bat material, Mr Henk Godthelp for making
available comparative fossil material and assisting with identification of rodents and Mr Ernst Holland for
advice and information about the Sooty Owl and geomorphology of Nettle Cave. Thanks also to Drs Alex
Baynes, Rod Wells and Suzanne Hand for critically reading earlier drafts of this paper. The Jenolan Caves
Scientific Advisory Committee gave permission for the excavation in Nettle Cave.
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Proc. LINN. SOc. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 157
APPENDIX A
A list of the published descriptions used in identifying the non-volant, native mammal species from the Nettle Cave
deposit. The illustrated key to Australian Mammalia (Jones and Baynes 1989) was used to check all specimens.
Taxon Reference
Antechinus Davison (1986); Merrilees & Porter (1979); Van Dyck (1982);
Wakefield & Warneke (1967)
Sminthopsis Archer (1981); Merrilees & Porter (1979)
Phascogale Merrilees & Porter (1979)
Dasyurus Green (1983); Merrilees & Porter (1979)
Isoodon, Perameles
Pseudocheirus
Petaurus
Petauroides
Cercartetus
Burramys parvus
Acrobates pygmaeus
Potorous
Bettongia
Thylogale thetis
Rattus
Pseudomys
Mastacomys
Freedman (1967); Freedman & Joffe (1967); Freedman & Rightmire (1971);
Green (1983); Lyne & Mort (1981); Merrilees (1967); Merrilees & Porter (1979)
Green (1983); Merrilees & Porter (1979)
Green (1983)
Archer (1984)
Green (1983); Merrilees & Porter (1979); Turnbull & Schram (1973)
Dixon (1971)
Archer (1984)
Green (1983); Merrilees & Porter (1979)
Green (1983); Merrilees & Porter (1979); Wakefield (1967)
Green (1983)
Green (1983); Merrilees & Porter (1979); Musser (1981); Tate (1951);
Taylor & Horner (1973)
Green (1983); Merrilees & Porter (1979); Schram & Turnbull (1970)
Green (1983)
Proc. LINN. Soc. N.S.W., 117. 1997
158 LATE QUATERNARY DEPOSIT
APPENDIX B
The minimum number of individuals (MNJ) of small mammals and skinks identified from the Nettle Cave deposit. Birds and agamids |
are listed as present (yes) from the depth intervals of the deposit in which they were found. Depth intervals are in cm. The relative |
abundance (%) and aggregate (AGG), i.e. the total number of specimens of the mammal species in each depth interval is also given. |
L1, L2, etc. represents levels 1, 2 and so on. Depth intervals 41-39 cm and 13-5 cm were not analysed. 1. These do not represent new |
species in the relevant genus; the specimens were broken or lacking teeth, thus could not be assigned to a species with certainty. 2. If |
generic identification is correct, the species would be M. schreibersii. 3. If generic identification is correct, the species would be
C. gouldii.
Depth interval 68-66 (L16) 66-64 (15) 64-59 (L14) 59-54 (L13) 54-50 (L12) 50-46 (L11) 46-44 (L10) L
Species MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%)AGG MNI(%) AGG) (i)
DASYURIDAE 1 (9.1) 1 2 (14.3) 2 3 (10.7) 3 3 (9.1) 4 3 (8.1) 6 7 (16.7) 9 8 (10.3) 12 | 5
Antechinus stuartii sensu lato 0 0 0 0 0 0 1 (1.3) 1 i
A. swainsonii 0 1(7.1) 1 1 (3.6) 1 2 (6.1) 3 2 (5.4) 4 2 (4.8) 4 1 (1.3) 1 i
A. flavipes 0 0 0 0 0 0 2 (2.6) 4 !
A. spp.' 0 0) 0 1(3)1 0 3 (7.1) 3 2 (2.6) 3 |
Sminthopsis murina 0 0 1 (3.6) 1 0 1 (2.7) 2 0 1 (1.3) 2
S. sp.’ (0) 0 0 0 0 1 (2.4) 1 0
Phascogale tapoatafa 0 0 1 (3.6) 1 0 0 0 1 (1.3) 1
Dasyurus sp. cf. D. viverrinus 0 1(7.1) 1 0) 0 0 1 (2.4) 1 0
a NT
PERAMELIDAE 1 (9.1) 1 0 2 (7.1) 2 1(3)1 1 (2.7) 1 1 (2.4) 1 1 (1.3) 1 ’
Isoodon obesulus 0 0 1 (3.6) 1 0 0 0 0
Perameles nasuta 1 (9.1) 1 0 1 (3.6) 1 1@)1 1 (2.7) 1 1 (2.4) 1 1(1.3) 1 if
PSEUDOCHEIRIDAE 0 1 (7.1) 1 0 0 0 1 (2.4) 1 0
Pseudocheirus peregrinus 0 1 @a) i 0 0 0 1(2.4) 1 0 r
Petauroides volans 0 0 0 0 0 0 0
PETAURIDAE 1(9.1) 1 0 0 0 1 (2.7) 1 0 0
Petaurus breviceps 1(9.1) 1 0 0 0 1 (2.7) 1 0 0 :
BURRAMYIDAE 0 2 (14.3) 3 2 (7.1) 3 4 (42.1) 7 4 (10.8) 7 8 (19) 18 14 (17.9) 33 |
Cercartetus nanus 0 0 0 0 0 0 0 4
C. lepidus 0 0 1 (3.6) 1 1 (3) 1 0 0 3 (3.8) 3
Burramys parvus 0 2 (14.3) 3 1 (3.6) 2 3 (9.1) 6 4 (10.8) 7 8 (19) 18 11 (14.1) 30 |
ACROBATIDAE 0 0 0 0 0 0 0
Acrobates pygmaeus 0 0 0 0 0 0 0
POTOROIDAE 0 0 0 0 0 0 10.3)1 §
Potorous sp. cf. P. tridactylus 0 0 0 0 0 0 13) ae
Bettongia sp. 0 0 0 0 0 0 0
MACROPODIDAE 0 0 0 0 0 0 0
Thylogale thetis 0 0 0 0 0 0 0
MURIDAE 8 (72.7) 14 9 (64.3) 19 21 (75) 46 25 (75.8) 60 28 (75.7) 62 24 (57.1) 53 52 (66.7) 138 |
Conilurus albipes 0 0 0 0 0 0 0
Pseudomys oralis 1 (9.1) 1 Gai) 72 4 (14.3) 7 3 (9.1) 8 3 (8.1) 7 3 (7.1) 4 6 (7.7) 16
P. gracilicaudatus 0 0 0 0 0 0 0
P. australis 0 (a3) 3) 1 (3.6) | 2 (6.1) 3 2 (5.4) 2 0 2 (2.6) 4
P. novaehollandiae 0 0) 0 0 0 0 0)
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD 159
44-43 (L9) 43-41 (L8) 39-37 (L6) 37-35 (L5) 35-25 (L4) 25-13 (L3) 5-0 (L1) SURFACE
MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%) AGG MNI(%) AGG = MNI(%) AGG
6 (16.7) 10 14 (13.6) 33 51 (13.6) 115 92 (20.3) 198 175 (19.1) 324 90 (12.4) 197 235 (16.6) 564 13 (10.8) 32
0 3 (2.9) 5 16 (4.3) 38 17 (3.8) 40 26 (2.8) 48 20 (2.8) 51 74 (5.2) 202 5 (4.2) 17
0 0 2 (0.5) 2 3 (0.7) 3 10 (1.1) 12 13 (1.8) 19 18 (1.3) 30 0
2 (5.6) 3 3 (2.9) 7 9 (2.4) 18 7 Gl) Ns) 15 (1.6) 28 18 (2.5) 41 45 (3.2) 101 2 (1.7) 3
2 (5.6) 3 0 0 23) (9.1) 55 45 (4.9) 78 18 (2.5) 35 50 (3.5) 118 4 (3.3) 10
2 (5.6) 4 8 (7.8) 21 22 (5.9) 55 27 (6) 63 46 (5) 114 15 (2.1) 43 31 (2.2) 86 2 (1.7) 2
0 0 0 12 (2.6) 19 17 (1.9) 26 3 (0.4) 4 10 (0.7) 15 0
0 2 (0.5) 2 2 (0.4) 2 10 (1.1) 12 3 (0.4) 4 6 (0.4) 11 0
0 0 0 1 (0.2) 1 6 (0.7) 6 0 1(0.1) 1 0
1 (2.8) 1 3 (2.9) 3 11 (3) 16 22 (4.8) 27 31 (3.4) 45 21 (2.9) 38 34 (2.4) 43 1 (0.8) 1
1 (2.8) 1 1()1 4 (1.1) 4 7 (1.5) 8 14 (1.5) 20 3 (0.4) 3 16 (1.1) 23 0
0 2 (1.9) 2 7 (1.9) 12 15 (3.3) 19 17 (1.9) 25 18 (2.5) 35 18 (1.3) 20 1 (0.8) 1
2 (5.6) 3 5 (4.9) 5 8 (2.2) 12 13 (2.9) 33 15 (1.6) 31 26 (3.6) 65 62 (4.4) 136 21 (17.5) 56
2 (5.6) 3 4 (3.9) 4 7 (1.9) 11 13 (2.9) 33 13 (1.4) 29 20 (2.8) 58 54 (3.8) 127 19 (15.8) 54
0 1()1 1 (0.3) 1 0 2 (0.2) 2 6 (0.8) 7 8 (0.6) 9 2 (1.7) 2
0 4 (3.9) 8 12 (3.2) 24 19 (4.2) 44 26 (2.8) 60 33 (4.6) 70 107 (7.6) 306 43 (35.8) 136
0 4 (3.9) 8 12 (3.2) 24 19 (4.2) 44 26 (2.8) 60 33 (4.6) 70 107 (7.6) 306 43 (35.8) 136
3 (8.3) 6 1(1)1 3 (0.8) 3 2 (0.4) 2 4 (0.4) 4 0 10 (0.7) 15 0
0 1(1)1 3 (0.8) 3 2 (0.4) 2 4 (0.4) 4 0 10 (0.7) 15 0
1 (2.8) 1 0 0 0 0 0 0 0
2 (5.6) 5 0 0 0 0 0 0 0
0 0 5 (1.3) 7 4 (0.9) 6 6 (0.7) 6 11 (0.8) 16
0 0 5 (1.3)7 4 (0.9) 6 6 (0.7) 6 0 11 (0.8) 16 0
0 0 4(1.1) 4 5 (1.1) 5 5 (0.5) 5 6 (0.8) 6 8 (0.6) 9 0
0 0 3 (0.8) 3 4 (0.9) 4 3 (0.3) 3 4 (0.6) 4 7 (0.5) 8 0
0 0 1 (0.3) 1 1 (0.2) 1 2 (0.2) 2 2 (0.3) 2 1 (0.1) 1 0
1 (2.8) 1 0 0 1 (0.2) 1 0 0 0 0
1 (2.8) 1 0 0 1 (0.2) 1 0 0 0 0
22 (61.1) 49 74 (71.9) 192 277 (73.8) 755-294 (64.9) 839 = 649. (71) 1833 541 (74.8) 1485 932 (65.8) 2920 40 (33.3) 83
0 1(1)1 2 (0.5) 2 6 (1.3) 7 23 (2.5) 57 2 (0.3) 2 1 (0.1) 1 Pd (ea) 22
3 (8.3) 8 17 (16.5) 44 80 (21.3) 253 98 (21.6) 320 236 (25.8) 740 = 256 (35.4) 860 465 (32.8) 1602 4 (3.3) 9
0 0 0 0 1 (0.1) 1 0 0 0
3 (2.9) 4 7 (1.9) 11 9 (2) 11 3 (0.3) 3 4 (0.6) 5 6 (0.4) 6 0
1 (2.8) 1 5 (4.9) 14 28 (7.5) 82 33 (7.3) 84 132 (14.4) 391 40 (5.5) 104 35 (2.5) 75 0
Proc. LINN. SOc. N.S.W., 117. 1997
160 LATE QUATERNARY DEPOSIT
:
Depth interval 68-66 (L16) 66-64 (15) 64-59 (L14) 59-54 (L13) 54-50 (L12) 50-46 (L11) 46-44 (L10) |
|
Species MNI(%) AGG MNI(%) AGG MNI(%)AGG MNI(%)AGG MNI(%)AGG MNI(%) AGG MNI(%) AGG
P. fumeus 0 1(7.1) 1 1 (3.6) 1 2 (6.1) 4 2 (5.4) 2 2 (4.8) 5 5 (6.4) 16
P. spp.' 2 (18.2) 2 1(7.1)4 1 (3.6) 1 2 (6.16 2 (5.4) 2 5 (11.9) 11 7 (9) 22
Mastacomys fuscus 5 (45.5) 10 4 (28.6) 8 13 (46.4) 35 15 (45.5) 38 15 (40.5) 39 11 (26.2) 28 25 (32.1) 62 |
Rattus fuscipes 0 1 (7.1) 3 1 (3.6) 1 0 4 (10.8) 6 2 (4.8) 4 5 (6.4) 15
R. rattus 0 0 0 0 0) 0) 0
R. spp." 0 0 0 1 (3) 1 0 1 (2.4) 1 2 (2.6) 3
Mus musculus 0 0 0 0 0 0 0
VESPERTILIONIDAE 0 0 0 0 0 1 (2.4) 1 2 (2.6) 2
Nyctophilus sp. cf. N. gouldi 0 0 0 0 0 0 1 (1.3) 1
N. sp. cf. N. geoffroyi 0 0 0 0 0 1 (2.4) 1 1 (1.3) 1
Miniopterus schreibersii 0 0 0 0 0 0 0
cf. Miniopterus* 0 0 0 0 0 0 0
Chalinolobus gouldii 0 0 0 0 0 0 0
C. morio 0 0 0 0 0 0 0
cf. Chalinolobus* 0 0 0 0 0 0 0
Falsistrellus tasmaniensis 0 0 0 0 0 0 0
LEPORIDAE 0 0 0 0 0 0 0
Oryctolagus cuniculus 0 0 0 0) 0 0 0
TOTAL MNI (%) 11 (100) 14 (100) 28 (99.9) 33 (100) 37 (100) 42 (100) 78 (100.1)
TOTAL AGGREGATE 17 26 54 72 i, 83 187
AVES yes yes yes yes yes yes yes
AGAMIDAE 0 0 0 0 0 0 0
SCINCIDAE 0 0 0 0 0 0 0
Egernia sp. cf. E. whitii 0 0 0 0 0 0 0
Eulamprus quoyii sp.-group 0 0 0 0 0 0 0
New taxon 0) 0 0 0) 0 0 0
Proc. LINN. SOC. N.S.W., 117. 1997
D.A. MORRIS, M.L. AUGEE, D. GILLIESON AND J. HEAD
161
44-43 (L9)
MNI(%) AGG = MNI(%) AGG
43-41 (L8)
39-37 (L6)
37-35 (L5)
35-25 (L4)
25-13 (L3)
MNI(%) AGG MNI(%)AGG MNI(%)AGG MNI(%) AGG
5-0 (L1)
SURFACE
MNI(%) AGG = MNI(%) AGG
2 (5.6) 4
4 (11.1) 10
6 (16.7) 16
6 (16.7) 10
0
0
0
1 (2.8) 1
0
1 (2.8) 1
Se So So 2Se CO ©
36 (100.1)
71
yes
8 (7.8) 17
13 (12.6) 35
10 (9.7) 24
16 (15.5) 51
0
1(1) 1
0
2 (1.9) 2
0
1()1
0
103 (100.1)
244
yes
14 (3.7) 32
35 (9.3) 89
25 (6.7) 63
63 (16.8) 184
0
23 (6.1) 39
0
4(1.1)4
0
2 (0.5) 2
0
0
1 (0.3) 1
0
0
1 (0.3) 1
0
0
375 (100.1)
940
yes
yes
0
0
0
0
15 (3.3) 34
25 (5.5) 71
22 (4.9) 57
68 (15) 220
0
18 (4) 35
0
1 (0.2) 1
Soo oS © © - ©
1 (0.2) 1
0
0
0
453 (99.9)
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54 (5.9) 145
31 (3.4) 69
118 (12.9) 339
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21 (2.9) 50
47 (6.5) 139
151 (20.9) 292
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311
Proc. LINN. Soc. N.S.W., 117. 1997
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Perameles bowensis, a New Species of
Perameles (Peramelemorphia, Marsupialia)
from Pliocene Faunas of Bow and
Wellington Caves, New South Wales
JEANETTE MUIRHEAD, LYNDALL DAWSON AND MICHAEL ARCHER
School of Biological Science, University of New South Wales, Sydney 2052
Muirhead, J., Dawson, L., and Archer, M. (1997). Perameles bowensis, a new species of
Perameles (Peramelemorphia, Marsupialia) from Pliocene faunas of Bow and
Wellington Caves, New South Wales. Proceedings of the Linnean Society of New
South Wales 117: 163-174
Perameles bowensis n. sp. is described from the early Pliocene Bow Local Fauna and
is phylogenetically compared to other species of Perameles. This species is the second
species of Tertiary Perameles. It appears to be closer to Recent species of Perameles than to
the early Pliocene Perameles allinghamensis. Specimens from Pliocene sediments of
Wellington Caves are also considered to represent P. bowensis. The presence of this species in
both the Bow and Big Sink Local Fauna (Wellington) provides further support to the correla-
tion of these faunas.
Manuscript received 17 June 1996, accepted for publication 23 October 1996.
KEYWORDS: Bandicoot, Perameles, Pliocene, Wellington Caves, Bow Local Fauna, Big
Sink Local Fauna
INTRODUCTION
Perameles allinghamensis Archer, 1976 was the first species described from the fossil
record of the otherwise modern genus Perameles. It was recovered from the early Pliocene
Bluff Downs Local Fauna in north Queensland (Archer 1976). Other Tertiary occurrences of
Perameles have been reported, e.g. the Miocene Tarkarooloo and Kutjamarpu Local Faunas
(Rich et al. 1982), however, none of these have been formally described and re-examination
of this material indicates that these specimens have been incorrectly attributed to Perameles
(Rich et al. 1991). Perameles bowensis from the Bow Local Fauna of northern New South
Wales is the second species of Tertiary Perameles described.
There are currently five recognised species of Perameles. These are P. nasuta
Geoffroy, 1804, P. gunnii Gray, 1838, P. bougainville Quoy and Gaimard, 1824, P. eremi-
ana Spencer, 1897 and the Pliocene P. allinghamensis Archer, 1976. While Tate (1948)
regards P. bougainville, P. eremiana, P. fasciata and P. notina to be possible local races of
a widespread southern species, Marlow (1962) considers all of these to be distinct species.
Here these forms are collectively regarded as P. bougainville except for P. eremiana which
is considered distinct following Mahoney and Ride (1988) and Seebeck et al. (1990). All
comparative specimens used in this work representing P. bougainville were collected as
subfossil specimens from the Nullarbor Plains in Western Australia and therefore do not
necessarily represent all possible races and variants of this species.
Dental nomenclature used follows that of Flower (1864) and Luckett (1993)
where the adult (unreduced) tooth formula of marsupials is P1-3 and M1-4. Tooth mor-
phology nomenclature follows Muirhead (1994). Classification follows Aplin and
Archer (1987). Catalogue number abbreviations are AMF; Australian Museum palaeon-
tological collection.
Proc. LINN. Soc. N.S.W., 117. 1997
A NEW SPECIES OF PERAMELES
Figure 1. Perameles bowensis type material. A and A’ = AMF98810 stereo occlusal views. B and B’ =
AMF98809 stereo occlusal views. C and C’ = AMF98811 stereo occlusal views. D = AMF98811 lateral view
gual side. Scale bar at A = Imm. Scale bar at B = n. Scale bar at C and D = Imm.
PENS OF)
J. MUIRHEAD, L. DAWSON AND M. ARCHER 165
SYSTEMATICS
Supercohort: Marsupialia (Illiger, 1811) Cuvier, 1817
Cohort: Australidelphia Szalay, 1982
Order: Peramelemorphia (Kirsch, 1968) Aplin and Archer, 1987
Family: Peramelidae (Gray, 1825), (sens. Groves and Flannery, 1990)
Perameles bowensis n. sp.
Holotype
AMF98809 (Fig. 1B) Right M>
Paratypes
AMF98810 (Fig. 1A) RM2; AMF98811 (Fig. 1C and D) RP?; AMF98812 (Fig.
2C and D) RM3; AMF98813 (Fig. 2A and B) LM}.
Specific etymology
The species name is in reference to the type locality.
Type locality and age
The Bow Local Fauna lies within unnamed roadcut sediments in northeastern New
South Wales. This fauna has been dated at early Pliocene based on biocorrelation with
the radiometrically dated Bluff Downs Local Fauna (Skilbeck 1980, Flannery and Archer
1984, Rich et al. 1991).
Diagnosis
Perameles bowensis differs from all other species of Perameles in the following
combination of features: 1) small size, 2) greater development of metastylar region on
M>~, 3) shallow ectoflexus on M” and none on M*, 4) small hypocone on M~’, 5) large
Lng displacement of stylar cusps on M*, 6) small development of parastylar region on
M~¢, 7) strongly curved preparacrista on M*, 8) anterobuccal cingulum on M~ and M
but not connecting to preprotocrista, 9) P° ovoid in shape with well developed posterior
heel not continuing anteriorly beyond lingual side of primary cusp, 10) no anterior cusp
on P~, and 11) posthypocristid on M, not continuous to posterolingual corner of crown.
Perameles bowensis is phenetically most similar to P. eremiana in terms of size
and general shape of the molars. It differs from P. eremiana in the following features: 1)
more continuously thick cingulum between lingual to posterior corners of P~, 2) less
well Tee IOROS and shallow trough between St B and St D and associated crests on M
3) larger M” formed by the wider stylar shelf region and larger metastylar corner, 4) sty-
lar cusps larger and wider on M~, 5) postmetacrista entirely straight and not curved pos-
terobuccally, 6) hypocone much smaller, 7) angle of posthypocristid orientated more
oblique to the tooth length and not perpendicular, and 8) paracristid and metacristid of
My, and M3 are more distant.
Description
P3 crown is short and ovoid in occlusal shape. The posterior dimension of the
crown is wider than the anterior. The primary cusp is anteriorly positioned. One major
crest extends posteriorly from the primary cusp to a small posterior cusp. Expansion of
the crown is present as a heel from the lingual side of the primary cusp to a posterior
cusp. Slight development of a heel is also present on the buccal side of the crest.
Proc. LINN. Soc. N.S.W., 117. 1997
A NEW SPECIES OF PERAMELES
igure 2. Perameles bowensis paratype material. A and A—AMF98813 stereo occlusal views. B = AMF98813
Jateral view. C = AMF98812 lateral view. I AMF98812 stereo occlusal views. All scale bars = Imm. C and
D to same s
J. MUIRHEAD, L. DAWSON AND M. ARCHER 167
M2 crown is roughly square in occlusal outline. The posterobuccal corner is bro-
ken. The anterior cingulum terminates below the midpoint of the preparacrista. No con-
tact exists between the anterior cingulum and the preprotocrista. The preprotocrista ter-
minates at the anterolingual corner of the paracone base. The crown is worn. The largest
cusp is St D, followed (in decreasing height) by St B, metacone, anterior cingulum tip,
paracone, protocone and hypocone. The preparacrista is short and curves to connect to
the anterior cingulum tip without connection to St B. The postparacrista is slightly short-
er in length, terminating around the posterior base of St B. The premetacrista is slightly
longer than the preparacrista and parallel to this crest, terminating at the anterolingual
base of St D. The postmetacrista 1s very worn and broken posterobuccally. A slight
inflection exists between the protocone and hypocone. The posthypocristid terminates at
the posterior base of the metacone. St B and D are the only stylar cusps present. These
have strong lingual curvature. The paracone is located midway between the buccal and
lingual sides of the crown. The metacone is positioned at a more lingual position. The
protocone and hypocone are equidistant from the buccal edge.
M? crown is triangular in occlusal view. The buccal surface is the shortest of the
three crown dimensions. All cusps and crests are worn. Morphology follows that of M<
except as follows. Ectoflexus is stronger on the buccal side. The posterobuccal corner of
the crown is prominently extended. St B and D do not curve lingually as much as on M°-.
The anterior cingulum is shorter. The position of the paracone is more lingual and lies
directly anterior to the metacone. The paracristae are longer. The postparacrista connects
directly to St B rather than to the base of this cusp. The trough between St B and D is
deeper at the shelf. The hypocone is small. The crest from the hypocone terminates at the
lingual base of the metacone. The parastylar region is developed to a greater degree, with
a larger distance between St B and the anterior cingulum tip.
The protoconid and metaconid of My are approximately equal in height, followed
(in decreasing height) by the entoconid, hypoconid, hypoconulid and paraconid. Of the
primary cusps, the metaconid and protoconid are closer to each other than either is to the
paraconid. The metaconid is directly posterior to the paraconid. The entoconid lies
directly posterior to the metaconid and not connected by a preentocristid. The entoconid
is conical in shape. The hypoconid is positioned almost twice as far buccally as is the
protoconid. The posthypocristid is the longest crest on the crown connecting to the
hypoconulid at the posterobuccal base of the entoconid. The cristid obliqua is curved,
terminating at the posterior base of the protoconid and buccal to the valley in the
metacristid. The entoconid lies directly lingual to the hypoconid. No anterior or posterior
cingulum is present.
Morphology of M3 follows that of Mj except as follows. The metaconid is higher
than the protoconid. The anterior cingulum is wide and without a notch, terminating at
the anterior base of the protoconid. The metaconid is closer to the paraconid than to the
protoconid. The protoconid is higher than the hypoconid. The protoconid is almost buc-
cally level with the hypoconid. The cristid obliqua terminates at the base of the valley in
the metacristid. No hypoconulid is present. The posthypocristid terminates at the base of
the entoconid. A wear facet along the crest of the entoconid lies oblique to the antero-
posterior length of the tooth.
Measurements of P. bowensis type material are found in Table 1.
PERAMELES BOWENSIS FROM WELLINGTON CAVES
Material collected from the Phosphate Mine Beds of Wellington Caves (NSW) dur-
ing excavations by A. Osborne, M. Archer and L. Dawson in 1982-1983, has produced
two bandicoot taxa (Dawson and Augee this volume). Five of these specimens appear to
be Perameles bowensis.
Proc. LINN. SOC. N.S.W., 117. 1997
168 A NEW SPECIES OF PERAMELES
All five specimens are isolated molars. Three specimens are from the Big Sink
Unit and include two broken Ms (AM F69887 [formerly WC1678], Fig 3B and AM
F69899, Fig 3A) and an BoM (AM F69804 [formerly WC1677], Fig 3C). The remaing
two specimens, a broken RM* (AM F69897, Fig 3E) and a RM* (AM F69896, Fig 3D),
were retrieved from the lower “Graded-Bedded Unit’ (Osborne 1982).
The *“Graded-Bedded Unit’ is separated from the Big Sink Unit by the discon-
formably overlying “Conglomerate Unit’ (Osborne 1983). The unconformity separating
the Big Sink Unit from the overlying Mitchell Cave Beds has been estimated by Osborne
(1983) to be at least late Pliocene in age. Teeth from both of these deposits appear to rep-
resent the same taxon.
TABLE 1
Measurements of specimens of Perameles bowensis from the Bow Local Fauna and Wellington Caves. All
measurements are maximum distances in mm. Width is lingual-buccal distance on crown. Length is antero-pos-
terior distance. Para = paracone, meta = metacone, proto = protocone, ento = entoconid, hypo = hypoconid,
metad = metaconid, parad = paraconid, protod = protoconid, — = information missing or not appropriate.
Specimens from Bow
Uppers
AME number width length para-meta meta-proto proto-para
98810 2.54 2.50 1.27 1.18 0.77
98809 2.76 3.14 1.53 1.74 0.77
98811 1.69 1.88 — — -
Lowers
AMF number width length ento-meta meta-hypo metad-parad parad-protod protod-metad
983813 1.61 2.52 0.92 1.12 1.02 1.07 0.69
98812 1.84 2.81 1.14 1.42 0.71 1.27 1.16
Specimens from Wellington Caves
Uppers
AMF number width length para-meta meta-proto proto-para
69896 2.83 2.61 1.63 1.79 0.95
69899 - 3),517/ 159 - -
69887 - - - - -
69897 - 3.34 1.78 — ~
Lowers
AMF number width length ento-meta meta-hypo metad-parad parad-protod protod-metad
69804 1.63 - 1.01 1.30 ~ - 0.79
These bandicoots specimens from Wellington Caves are clearly a Perameles rather
than an /soodon due to the lack of complete anterior and posterior cingulum on the upper
molars (particularly the posterior molars). These specimens are within the size range of
P. bougainville and P. eremiana; however they differ in morphology from these species.
The only remaining species of Perameles of this size range is P. bowensis from the Bow
Local Fauna,
No M!s are available in the Bow Local Fauna attributed to P. bowensis. This there-
Proc. LINN. SOc. N.S.W., 117. 1997
J. MUIRHEAD, L. DAWSON AND M. ARCHER
Figure 3. Perameles bowensis from Wellington Caves. Stereo occlusal views of specimens. A and A’ =
AMF69899. B and B’ = AMF69887. C and C’ = AMF67804. D and D’ = AMF69896. E and E’ = AMF69897.
Scale bars = 2mm. Lower scale bar refers to all specimens except A.
Proc. LINN. Soc. N.S.W., 117. 1997
170 A NEW SPECIES OF PERAMELES
fore prevents direct comparison between the M! from Wellington Caves and this species.
The morphology of the broken MI, however, is within the range exhibited by other
species of Perameles and within that expected following the morphological trends of the
more posterior molars of this species.
The M2 from Wellington Caves is very similar to the corresponding tooth from P.
bowensis in having no ectoflexus, no anterior cingulum, the same development of the
posterior cingulum, the same degree of lingual displacement of the stylar cusps and a
similar height of all of the cusps on the tooth. The metastylar region cannot be compared
because this area has been broken in the Bow sample. The Wellington M~ differs from
the corresponding tooth from Bow in having a less concave preparacrista and greater dis-
similarity in size between the protocone and hypocone. This difference in morphology
between the M* from these sites is easily attributed to wear. The M* from Bow shows
greater wear at the back of the lingual tip of the protocone. These is little reason to doubt
the inclusion of this tooth within the morphological and size range expected for P.
bowensis.
The Ms from Wellington Caves are represented by two broken fragments. The
larger of these includes the paracone, metacone and the stylar region of this tooth. The
smaller fragment represents the paracone and parastylar region and in all respects dupli-
cates the morphology and size shown by the more completely preserved specimen. The
M?s from Wellington are similar to the corresponding tooth of P. bowensis in all respects
except for the lesser ectoflexus with less extension of the postmetacrista and the greater
size of the St E. The size difference of St E may be attributed to wear. The extent of mor-
phological variation for this species is unknown because of the very limited number of
samples, but this degree of dfference is unlikely to be beyond that exhibited within a
species.
The RM, differs from that of P. bowensis from the Bow Local Fauna only by a
slightly smaller hypoconid.
There is little to preclude specimens from the Big Sink Unit and the “Graded
Bedded Unit’ of Wellington Caves from being considered representative of P. bowensis.
All differences between specimens may be attributed to wear or intraspecific variation.
Because specimens from the type locality do not include an example of the M1! this
tooth of P. bowensis is described from Wellington Caves.
Description of mM!
The buccal portion of the M! is the only part preserved. The highest cusp on the
crown is the metacone, followed (in decreasing height) by the St D, St E, paracone and
St B. The parastylar region includes a small, unnotched tip on the anterobuccal corner of
the crown. The paracrista does not connect to the parastylar tip but instead runs poster-
obuccally from the paracone to connect with St B. The postparacrista runs parallel to the
preparacrista from the paracone to connect at the posterior flank of the St B. There is no
ectoflexus on the crown and the buccal face of the tooth is slightly rounded. The trough
between St B and St D is shallow. St D is conical in shape and there is no connection by
way of a crest to the St B. The premetacrista is straight and terminates at the anterior
base of St D. The postmetacrista is the longest crest on the tooth, terminating at the pos-
terobuccal metastylar tip of the crown. It is slightly convex around the metastylar region.
St E lies on the posterobuccal region of the stylar shelf. It has a short anterior and poste-
rior crest connecting to the St D and metastylar tip respectively. The broken region at the
base of the metastylar portion of the tooth indicates a triangular shape of the complete
tooth with the presence of a hypocone. The posterior cingulum would, if present, not
have connected to the base of the metastylar corner of the crown.
Proc. LINN. SOc. N.S.W., 117. 1997
J. MUIRHEAD, L. DAWSON AND M. ARCHER 171
PHYLOGENETIC DISCUSSION
Aplin and Archer (1985) recognise the presence of both Jsoodon and Perameles
within the Bow Local Fauna. Part of the material described here as Perameles bowensis
was that referred to as Jsoodon (Aplin and Archer 1986, Archer 1984, Rich et al. 1991).
There is no material from Bow that shares apomorphies with /soodon that are not also
shared with Perameles (e.g. enlargement of the hypocone on M2). Apomorphies that dis-
tinguish Jsoodon from Perameles are not apparent in this material such as the complete
(or almost complete) and well developed anterior and posterior cingula on M? and M?,
and enlarged roots with a lack of distinction between root and crown. These specimens
are therefore precluded from /soodon. The specimens previously considered to represent
Isoodon are now included with material referred to as Perameles and described here as
the one species, P. bowensis.
Material refered to here as Perameles bowensis shares no apomorphies with any
other genus not also shared with other species of Perameles. Features that P. bowensis
has in common with other genera that are not also shared with other species of
Perameles are symplesiomorphies. The morphology of P. bowensis appears to fall well
within the range of diversity exhibited by modern species of Perameles and 1s therefore
placed within this genus.
In general, P. bowensis appears to be slightly more plesiomorphic in most regards
than modern species of Perameles but is likely to be closer to these than to P. alling-
hamensis. Of the modern species, P. bougainville is considered to show more plesiomor-
phic features of the dentition (but not cranial characters). Perameles bougainville, unlike
P. eremiana,_P. nasuta and P. gunnii, retains the plesiomorphic characteristics of less
caniniform I~, more linear PY, smaller hypocones, less developed posterior cingulum,
less elongated snout, retention of the hypoconulid on M>, more equidistant paraconids,
metaconids, and protoconids and an incomplete anterior cingulum on M}.
Perameles bowensis is more plesiomorphic than all Recent species in having a
smaller hypocone on the M~, more equidistant paraconids, metaconids and protoconids
on Mj_3 with the paraconid and metaconid wider apart on M, than on Recent species.
Of the Recent species, P. bowensis is phenetically similar to P. eremiana in having simi-
lar sized P° and anol ats emote for the M”, which is slightly larger in P. bowensis. The
morphology of the P? and M2 does not differ between these two species except that the
metastylar region is smaller in P. bowensis and in this feature is more similar to P.
bougainville. The lower molars of P. eremiana are more apomorphic in their concave
cristid obliquas producing a narrower talonid. Perameles eremiana is also more apomor-
phic than P. bowensis in the overall smaller width of the trigonid basin produced by
reduction of the distance between the paraconid and metaconid and orientation of associ-
ated crests more perpendicularly to the long axis of the tooth row. Perameles bowensis 1s
plesiomorphic in this regard in having a wider trigonid which is also wider than in any
other Recent species. Perameles bowensis is also more plesiomorphic than P. eremiana
in having a smaller hypocone on the M”, a feature in which it is also more plesiomorphic
than all Recent species of Perameles.
Comparison to P. eremiana is restricted due to the few available samples of this
species and therefore intraspecific variation for P. eremiana cannot be adequately
assessed. However, variation to the degree needed to include P. bowensis into this
species is much wider than that known for any other species. No species is known to
vary the orientation of crests of the molars to such a degree. The variation between sam-
ples of P. eremiana and P. bowensis is therefore considered to be outside of that for a sin-
gle species, and P. bowensis is therefore separated from P. eremiana. The kind of varia-
tion between these two species in some characters (e.g. crest orientation, basin width)
represents the two extremes shown within the entire genus.
Perameles bowensis retains some of the plesiomorphies seen in P. bougainville.
These are the small hypocone on M2? and roughly equidistant paraconid, metaconid, and
Proc. LINN. Soc. N.S.W., 117. 1997
172 A NEW SPECIES OF PERAMELES
protoconids. The P3 is more apomorphic than that of P. bougainville, with posterior
ante