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Alcheringa: An Australasian Journal of Palaeontology
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Palaeontological excavation and taphonomic investigation of the late
Pleistocene fossil deposit in Grant Hall, Victoria Fossil Cave, Naracoorte,
South Australia
REBECCA A. FRASERa; RODERICK T. WELLSb
a School of Biological Sciences, Flinders University, South Australia b School of Biological Sciences,
Flinders University, Adelaide, South Australia
Online publication date: 01 February 2010
To cite this Article FRASER, REBECCA A. and WELLS, RODERICK T.(2006) 'Palaeontological excavation and taphonomic
investigation of the late Pleistocene fossil deposit in Grant Hall, Victoria Fossil Cave, Naracoorte, South Australia',
Alcheringa: An Australasian Journal of Palaeontology, 30: 2, 147 — 161
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ISBN 0 9757894 5 7/2006/15 $3.00 ©AAP
Palaeontological excavation and taphonomic investigation
of the late Pleistocene fossil deposit in Grant Hall, Victoria
Fossil Cave, Naracoorte, South Australia
REBECCA A. FRASER and RODERICK T. WELLS
FRASER, R.A. & WELLS, R.T., 2006. Palaeontological excavation and taphonomic
investigation of the late Pleistocene fossil deposit in Grant Hall, Victoria Fossil Cave,
Naracoorte, South Australia. Alcheringa Special Issue 1, 147-161. ISBN 0 9757894 5 7.
Grant Hall chamber in Victoria Fossil Cave, Naracoorte, South Australia, contains a late
Pleistocene faunal assemblage, dated at between 206 and 76 Ka. Taphonomic and faunal
analyses indicate that the predominant mode of accumulation was via a surface exposed
pitfall trap. An avian predator, such as Tyto alba, may have been responsible for the
accumulation of small mammal remains. The faunal assemblage is taxonomically diverse
containing at least 47 taxa. It includes many browsing species such as Wallabia bicolour
and the extinct Sthenurine kangaroos and Zygomaturus trilobus, as well as small mammals
that require trees and a thick understorey. The Grant Hall fauna thus indicates the
presence of densely vegetated woodland, interspersed with small patches of open and
thickly grassed areas in the proximal vicinity of the old cave entrance. The relative
abundances and species composition of the Macropodidae fauna in Grant Hall are
significantly different from other faunal assemblages found at Naracoorte. This study
has provided palaeoecological information for a time period not previously investigated
at the Naracoorte Caves; detailed surveying of the chamber was undertaken as part of
the study.
Rebecca A. Fraser [r.fraser@swansea.ac.uk], School of Biological Sciences, Flinders
University, South Australia (current address Geography Department, University of Wales,
Singleton Park, Swansea) & Roderick T. Wells [rod.wells@flinder s.edu.au], School of
Biological Sciences, Flinders University, GPO Box 2100 Adelaide, South Australia;
received 4.8.2005, accepted 21.8.2006.
Key words: Naracoorte, Victoria Fossil Cave, Grant Hall, Late Pleistocene, taphonomy,
fossil assemblage, pitfall trap.
NARACOORTE CAVES contain a diverse range
of well preserved fossil faunas from late
Quaternary to present day. Fossil Chamber in
Victoria Fossil Cave has undergone nearly 30
years of excavation and has yielded a large and
diverse faunal assemblage (Wells et al. 1984).
Extensive palaeontological investigations have
been undertaken in Cathedral Cave (Brown &
Wells 2000), Wet Cave and Robinsons Cave
(McDowell 2001), and there have been many new
fossil cave discoveries in the local region (Reed
& Bourne 2000). The ages of a small collection
of cave deposits have recently been determined
with uranium-series dating of associated
speleothems (Ayliffe et al. 1998, Moriarty et al.
2000) and electron spin resonance (ESR) dating
of tooth enamel (Grünet al. 2001). This has
enabled these faunal assemblages to be placed
within a chronological framework.
Grant Hall chamber is located within the
Victoria Fossil Cave, approximately 50m from the
main Fossil Chamber; however, the fossil
deposits are entirely separate. The age of the
Grant Hall fossil deposit has been constrained
between 206 Ka and 76 Ka by uranium-series
dating of associated speleothems (Ayliffe &
Veeh 1988). The fossil-bearing sediments were
sandwiched between separate flowstones that
covered the limestone base of the cave and
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ALCHERINGA
148 FRASER AND WELLS
overlaid the sediment’s surface; most of the top
flowstone had been removed during prior
excavations (Ayliffe & Veeh 1988). In addition,
ESR dates on Zygomaturus tooth enamel,
sampled at a depth of approximately 60 cm from
the surface, yielded an age of 125 Ka (Grünet al.
2001).
The primary objective of this paper is to
present the palaeontological and taphonomic
findings of the 2000 excavation in Grant Hall.
Comparisons with other dated fossil deposits at
Naracoorte are also made, and in light of the
faunal composition and regional palaeoclimatic
evidence, a preliminary reconstruction of the
proximal palaeoenvironment is presented.
Methods and materials
Surveying and palaeontological excavation
Prior to commencing the palaeontological
excavation, the main topographical features of
the Grant Hall chamber were mapped. The top of
a stalagmite at the southern end of the chamber
was designated the primary datum point for all
surveying. A network of 1 m2 string-line grids
was pegged out over the main sediment covered
floor. The heights of the chamber floor and ceiling
relative to datum and the location of surface
features, such as stalagmites and limestone roof-
fall blocks, were surveyed at 50 cm intervals using
a tape measure, a hand held laser distance meter
and a laser dumpy level. The incline of the
surrounding rock-pile slopes, although irregular,
was measured with a clinometer and compass.
The depths of the sub-surface topography were
measured by probing the sediment floor with a 1
m long 10 mm diameter metal rod at 1 m2 intervals
along the string-line grid.
The palaeontological excavation was
located as close as possible to the speleothems
that were uranium-series dated in 1988 (Ayliffe
& Veeh 1988) and 1996 (Moriarty et al. 2000);
this ensured the fossils from this study were in
stratigraphic context with the dating results. Four
1 m2 square grids were excavated in 10 cm levels
using trowels, dental picks and brushes. Large
fossil bones were reliefed out of the surrounding
sediment. Their in situ positions were drawn on
a 10 cm grid sheet, photographed and measured
relative to datum. The strike and dip angles (to
the proximal end) of long bones (>10 cm) were
recorded. All fossils were given a unique
specimen number and bagged. At the laboratory,
the bones were cleaned of sediments, dried at
ambient temperature and preserved with
Mowital™. The surrounding sediments from
each spit were collected separately in large plastic
bags and removed from the cave. These were
wet sieved through 2 mm2 wire mesh to retrieve
small bones.
A 2 kg sample of sediment was taken from
each 10 cm level. The sediments were analysed
using hand specimens and the simple
gravitational settling technique of Day (1965)
was used to estimate particle size, grain shape,
sorting and clay content. The position of
limestone gravels, calcite fragments and calcrete
rocks were included in the sedimentary analysis,
and clasts larger than 10 cm in diameter were
marked on the excavation grid sheets.
Taxonomy and taphonomy
Each fossil specimen was identified to at least
genus and, where possible, species. Taxonomic
identifications were made using reference
skeletons and existing fossil collections held at
the Flinders University palaeontology
laboratory. Quantitative analysis included a tally
of the number of identifiable specimens (NISP),
and calculations of the minimum number of
individuals (MNI), species richness and the
relative abundances of species and skeletal
elements per stratigraphic layer. MNI is a derived
quantity representing the minimum number of
individuals necessary to account for all of the
skeletal elements of a particular taxonomic group
(Grayson 1984), and in this study it was based
on the most abundant element of each taxon.
MNI calculations accounted for age and size of
specimens, but left and right pairs were not
formally matched. The relative abundances of
species and element types were calculated as a
Fig.1 Plan view of the sediment floor of Grant Hall
Fossil Chamber. The transect through A-A is shown in
Fig 2.
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ALCHERINGA 149LATE PLEISTOCENE OF GRANT CAVE, NARACOORTE
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ALCHERINGA
150 FRASER AND WELLS
percentage of the total faunal sample of each
layer. In a mammalian skeleton not all elements
occur in equal frequencies, therefore the
following calculation of Andrews (1990) was
used to determine the relative abundances of
elements (Ri): Ri= (Ni/MNI x Ei) x 100,
where Ni is the quantity of elements in the
sample and Ei is the quantity of that element in
the particular animal’s skeleton.
The taphonomic analysis examined each
fossil bone for surface modifications, such as:
abrasions, predator tooth marks, root or acid
etching, calcite covering, levels of completeness,
breakage patterns and weathering. Breakage
patterns followed the descriptions of Marshall
(in Lyman 1994, p.139) and were categorised as:
columnar, crenulated, spiral, V-shaped,
compression, smooth or irregular perpendicular
and depressed. The weathering stages 1 to 5
followed those of Behrensmeyer (1978).
Results
Surveying and excavation
Fig. 1 shows a plan view of the Grant Hall chamber.
Fig. 2 shows the cross-section view through the
chamber at the point A-A on Fig 1.
It appeared there were two possible entrance
holes for the influx of sediment and bones. The
first was at the northwest end of the chamber:
there a path of dark red/brown sediment began
near the cave roof and flowed through a steep
rock pile directly onto the floor at the northwest
end of the chamber. The second was at the
western edge of the chamber: there a path of
pale sandy sediment merged through the base
of the rock pile and met the cave floor at the
southern end of the chamber. Surveying
demonstrated that sediment from the western
entrance did not flow onto the main sediment
floor or contribute material to the fossil deposit.
The most likely site where the sediment and
Fig. 2. Diagrammatic cross section through Grant Hall Fossil Chamber (points A-A as on Fig. 1).
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ALCHERINGA 151LATE PLEISTOCENE OF GRANT CAVE, NARACOORTE
Table 1. Summary of large mammal species per stratigraphic layer. * denotes an extinct species. † the NISP and MNI
counts for Macropus sp. indet and Sthenurus sp. indet were based on post cranial elements that could not be identified
past genus.
Layer 1 Layer 2 Layer 3 Total
TAXONOMIC GROUP NIS
PMNI NIS
PMNI NIS
PMNI NIS
PMNI
Macropus rufogriseus 6 3 24 10 20 9 50 22
Macropus sp. indet.†144 9 418 16 357 16 919 41
Protemnodon cf. brehus* 1 1 0 0 0 0 1 1
Wallabia bicolour 3 2 27 12 13 9 43 23
TOTAL SUBFAMILY
MACROPODINAE 154 15 469 38 390 34 101
387
Simosthenurus gilli* 8 2 27 6 14 3 49 11
Simosthenurus
occidentalis* 1 1 4 1 0 0 5 2
Procoptodon browneorum*0 0 1 1 0 0 1 1
Simosthenurus maddocki* 0 0 1 1 2 2 3 3
Sthenurus andersoni* 1 1 0 0 0 0 1 1
Sthenurus sp. indet.* † 21 2 167 8 123 4 311 14
TOTAL SUBFAMILY
STHENURINAE 31 6 200 17 139 9 370 32
TOTAL
MACROPODIDAE 185 21 669 55 529 43 138
3119
Thylacinus cynocephalus* 4 1 7 1 9 1 20 3
TOTAL THYLACINIDAE 4 1 7 1 9 1 20 3
Thylacoleo carnifex* 8 1 15 1 7 1 30 3
TOTAL
THYLACOLEONIDAE 8 1 15 1 7 1 30 3
Vombatus urinus 2 1 6 1 2 1 10 3
TOTAL VOMBATIDAE 2 1 6 1 2 1 10 3
Zygomaturus trilobus* 1 1 28 2 11 1 40 4
TOTAL
DIPROTODONTIDAE 1 1 28 2 11 1 40 4
Tachyglossus aculeatus 0 0 0 0 2 1 2 1
TOTAL
TACHYGLOSSIDAE 0 0 0 0 2 1 2 1
Phascolarctos cinereus 1 1 7 1 0 0 8 2
TOTAL
PHASCOLARCTIDAE 1 1 7 1 0 0 8 2
TOTALS FOR EACH
LAYER 201 26 732 61 560 48 149
3135
* denotes an extinct species. † the NISP and MNI counts for Macropus sp. indet and Sthenurus
sp. indet were based on post cranial elements that could not be identified past genus
1
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ALCHERINGA
152 FRASER AND WELLS
Table 2. Summary of small mammal species per stratigraphic layer.
Layer 1 Layer 2 Layer 3 Total
TAXONOMIC GROUP NISP MNI NISP MNI NISP MNI NISP MNI
Mammalia- Marsupalia
Cercartetus nanus 0 0 1 1 21 13 22 14
TOTAL BURRAMYIDAE 0 0 1 1 21 13 22 14
Antechinus flavipes 1 1 10 6 23 12 34 19
Antechinus sp. cf. A. flavipes 0 0 1 1 10 9 11 10
Dasyurus viverrinus 1 1 4 3 2 1 7 5
Phascogale tapoatafa 0 0 1 1 4 3 5 4
Sminthopsis crassicaudata 2 2 17 8 45 27 64 37
Sminthopsis murina 0 0 4 2 15 10 19 12
Sminthopsis sp. Indet. 0 0 0 0 50 30 50 30
TOTAL DASYURIDAE 4 4 37 21 149 92 190 117
Perameles sp. cf. gunnii 4 4 0 0 0 0 4 4
Perameles gunnii 2 1 6 4 31 16 39 21
Perameles bougainville 0 0 0 0 42 26 42 26
TOTAL PERAMELIDAE 6 5 6 4 73 42 85 51
Petaurus breviceps 0 0 1 1 3 2 4 3
TOTAL PETAURIDAE 0 0 1 1 3 2 4 3
Potorus platyops 3 2 5 5 5 3 13 10
Potorus tridactylus 0 0 32 14 25 14 57 28
Potorus sp. indet. 6 2 1 1 4 2 11 5
TOTAL POTOROIDAE 9 4 38 20 34 19 81 43
Pseudocheirus peregrinus 0 0 3 2 6 4 9 6
TOTAL PSEUDOCHERIDAE 0 0 3 2 6 4 9 6
Total Subclass Marsupalia 19 13 86 49 286 172 391 234
Mammalia- Placentalia
Mastacomys fuscus 0 0 0 0 2 1 2 1
Notomys mitchelli 3 1 6 4 11 7 20 12
Pseudomys apodemoides 0 0 14 6 93 43 107 49
Pseudomys australis 3 2 12 7 13 5 28 14
Pseudomys basalticus 0 0 2 1 4 2 6 3
Pseudomys shortridgei 6 3 0 0 57 26 63 29
Pseudomys sp. cf. P fumeus 0 0 17 6 34 10 51 16
Rattus fuscipes 3 2 34 10 67 20 104 32
Rattus lutreolus 0 0 3 1 2 1 5 2
Rattus tunneyi 1 1 6 4 18 9 25 14
Hydromys chrysogaster 0 0 0 0 1 1 1 1
TOTAL MURIDAE 16 9 94 39 302 125 412 173
Miniopterus sp. cf. schreibersii 0 0 1 1 0 0 1 1
TOTAL VESPERLLONIDAE 0 0 1 1 0 0 1 1
Total Subclass Placentalia 16 9 95 40 302 125 413 174
TOTAL FOR EACH LAYER 35 22 181 89 588 297 804 408
2
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ALCHERINGA 153LATE PLEISTOCENE OF GRANT CAVE, NARACOORTE
bones entered the cave was a hole at the
northwest end of the chamber. This is shown on
Fig. 1. The depths of the cave floor sediments
increased from negligible at the base of the rock-
pile to over 1 m at the eastern sidewall. The
deposit may go deeper, but the low ceiling there
prevented a longer probe being used.
The palaeontological excavation occurred
in the grids numbered 1, 2, 5 and 6, shown on
Fig. 1, from the surface to the speleothem-covered
limestone base. It is unknown whether this is
the cave floor or if more sediment layers exist
underneath. The maximum depth reached was
90 cm in Grid 6.
Sediments and stratigraphy
The sediments were largely homogeneous,
consisting of a mixture of red/brown and pale
yellow medium to fine sub-rounded quartz
grains, mixed with darker red-brown clay
particles. Despite the overall homogeneity of the
sediments, the fossil deposit was divided in to
three stratigraphic layers separated by two sub-
layers. Layers 1 (0-20 cm depth) and 2 (20-50 cm)
were separated by a 2-3 cm band of pale sediment
comprising a large proportion of gravel, and
pebble sized limestone clasts. Layers 2 and 3
(depth 50-90 cm)were separated by a 3-4 cm band
of pale yellow/white medium sized, sub-rounded,
moderately sorted quartz sand and intermixed
with gravel and pebble sized limestone clasts.
Clay content, from the sediment settling analysis,
for Layers 1, 2 and 3 was approximately 12, 16
and 15% respectively.
Palaeontology
Relative species abundances
Tables 1, 2 and 3 show the NISP and MNI counts
of each species identified in the Grant Hall fossil
assemblage. A total NISP of 2686 vertebrate
fossils were retrieved. The Macropodidae was
the most abundant large mammal group in the
deposit (MNI 119). Five species were identified
in the extinct Sthenurinae subfamily- the smaller
Simosthenurus gilli (Merrilees, 1965) appeared
in all three layers, and the larger sthenurines
occurred sporadically throughout the deposit.
Macropodinae subfamily was dominated by the
smaller kangaroos, Macropus rufogriseus
(Desmarest, 1817) and Wallabia bicolour
Table 3. Summary of reptile and avian species per stratigraphic layer. The majority of elements could only be
identified to genus level; thus MNI values were not calculated
TAXONOMIC GROUP Layer 1 NISP Layer 2 NISP Layer 3 NISP
Reptiles
Elapidae Gen. Sp. indet. 1 35 156
Wonambi naracoortensis* 0 0 3
Chelonia sp. Indet 0 3 1
Varanus sp. indet. 1 61 40
Pogona sp. cf. P. barbata
“Amphibolurus barbatus,
0 1 0
Tiliqua nigrolutea 0 1 21
Tiliqua rugosa 0 2 0
Scincidae Gen. Sp. indet. 0 0 2
TOTAL REPTILES 2 103 223
Aves 0 0 0
Aves Gen. Sp. indet. 0 0 16
TOTAL AVES 0 1 16
TOTAL NISP 2 104 237
3
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ALCHERINGA
154 FRASER AND WELLS
(Desmarest, 1804).A single molar was identified
as the extinct Protemnodon sp. cf. P. brehus
(Owen, 1874).
No specimens of the larger macropods
Macropus giganteus (Shaw, 1790) or M.
fuliginosus (Desmarest, 1817) were identified
from cranial-dental remains. Because these
species are extremely common in the other fossil
deposits at the Naracoorte Caves (Wells et al.
1984; Brown & Wells 2000) a closer examination
of the Macropus post-cranial elements was
undertaken to help clarify their absence. The
average length of specimens of the major limb
elements of the Grant Hall Macropus sp. indet.
was significantly smaller (~38%) than those of
Macropus giganteus measured by Flannery
(1981). Comparative measurements are shown
in Table 4. On this basis, it was concluded that
the post-cranial elements were most likely from
the smaller kangaroos (M. rufogriseus and W.
bicolour), rather than the larger Macropus
species; and thus, the later might be absent from
the faunal assemblage. However, the likelihood
that some of these elements came from juveniles
of the larger Macropus species cannot be
excluded.
The other herbivore taxa, Phascolarctos
cinereus (Goldfuss, 1817), Vombatus ursinus
(Shaw, 1800), and Zygomaturus trilobus
(Macleay, 1858) were relatively rare in the
deposit (MNI were 1 or 2 per layer). The
carnivore niche was represented by Thylacoleo
carnifex (Owen, 1859) and Thylacinus
cynocephalus (Harris, 1808). The small mammal
assemblage comprised 24 species; the greatest
species diversity occurred in the Dasyuridae and
the Muridae. Varanid and elapid vertebrae were
very common and avian remains were very rare.
The extinct giant snake, Wonambi naracoortensis
(Smith, 1976),was found only in Layer 3.
Taphonomy
Long bone orientation measurements did not
show any preferred alignments or dip angles.
Bones were patchily distributed laterally over
the grids, and bone concentration (per unit
volume of sediment) increased with excavation
depth. Dense concentrations of bone usually
contained a jumbled mix of skeletal elements of
varying sizes, whereas the sparser
concentrations contained mostly small elements
such as single phalanges, ribs, loose teeth and
vertebrae. Articulated and associated skeletal
elements were more often found near, or within,
the denser bone concentrations than in the
sparser patches. Nine sets of bone articulations
were found: the majority were between a) distal
limb elements like calcanei, astragali, cuboids and
metatarsals, and b) vertebrae.
Fig. 3 shows the results of the taphonomic
analyses of bone completeness categories,
breakage, weathering stages and surface
modifications, for each layer. The taphonomic
signatures were generally similar for each layer.
Overall, approximately 62% of all fossils were
complete. Table 5 shows the relative abundances
skeletal elements of all large mammal species in
each layer. Distal limb elements, such as
Table 4. Comparison of Macropus sp. indet.; limb measurements.
Average length in cm ± standard deviation Skeletal element
Grant Hall Macropus giganteus
after Flannery (1981)
% difference
Humeri 8.9 ± 1.3 14.1 36.9%
Ulnae 12.5 ± 2.1 19.8 36.9%
Radii 12.7 ± 0.6 18.1 29.8%
Femora 13.96 ± 3.8 24.8 44.0%
Tibia 24.1 ± 3.3 43.85 45.1%
4
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ALCHERINGA 155LATE PLEISTOCENE OF GRANT CAVE, NARACOORTE
Fig. 3. Results of taphonomic analysis: bone completeness, breakages and surface modifications.
metatarsals and tibiae, as well as pelvises and
dentaries, were the most abundant element
groups. Layer 1 had a significantly lower average
Ri (11%) than Layers 2 and 3 (20.9 and 22.9%,
respectively).
Discussion
Mode of accumulation
The quality of bone preservation in the Grant
Hall fossil assemblage was high. The low levels
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ALCHERINGA
156 FRASER AND WELLS
significant. There was little evidence for long
bone alignment due to continuous or strong
water flow, and bones lay relatively horizontal
with the stratigraphy. This suggested that once
animals were inside the cave and the carcasses
had disintegrated, their bones did not undergo
extensive reworking due to water currents or
sediment slumping. The low Ri values for the
small and light skeletal elements, such as
vertebrae, phalanges-manus and metacarpals,
might have been due to selective transport and
Table 5. Relative abundance (Ri) percentages; skeletal elements of all large mammal taxa.
of surface weathering and cracking suggested
that bones were deposited within the cave. The
natural trapping of unwary animals via a pit-fall
was therefore proposed as the predominant
method by which the large mammal bones came
to be in the cave.
The frequencies of taphonomic features
were similar for all layers (Fig. 3), which suggested
the modes of accumulation were similar through
time. Articulated and associated specimens were
rare, yet their presence at all was highly
Layer 1 Layer 2 Layer 3
Skeletal element NISP %Ri NISP %Ri NISP %Ri
skulls and cranial fragments 0 0 12 32.4 6 21.4
dentaries 9 24 42 56.8 25 44.6
maxillae 2 5 16 21.6 11 19.6
scapulae 1 3 9 12.2 8 14.3
clavicles 0 0 2 2.7 3 5.4
humeri 3 8 13 17.6 12 21.4
ulnae 3 8 16 21.6 12 21.4
radii 3 8 10 13.5 7 12.5
prox. phalanges - manus 0 0 1 0.3 0 0.0
medial phalanges - manus 6 3 14 3.8 7 2.5
distal phalanges - manus 1 1 6 1.6 1 0.4
pelvises 7 18 25 33.8 26 46.4
epipubic bones 0 0 4 10.8 4 14.3
femora 8 21 23 31.1 26 46.4
tibiae 8 21 34 45.9 38 67.9
fibulae 2 5 15 20.3 14 25.0
metatarsals (4th) 11 58 25 67.6 18 64.3
metatarsals (5th) 4 21 14 37.8 16 57.1
metacarpals 1 1 2 0.5 3 1.1
epiphysis of pubic symphysis 2 11 7.0 18.9 5 17.9
epiphysis of tibiae- proximal 3 8 12 16.2 12 21.4
epiphysis of tibiae- distal 4 11 12 16.2 13 23.2
epiphysis of femora- proximal 1 3 6 8.1 2 3.6
epiphysis of femora- distal 0 0 9 12.2 7 12.5
cuboids 3 8 9 12.2 9 16.1
calcanei 8 21 20 27.0 15 26.8
astragali 9 24 16 21.6 6 10.7
AVERAGE Ri for Layer 11 20.9 22.9
5
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ALCHERINGA 157LATE PLEISTOCENE OF GRANT CAVE, NARACOORTE
sorting under weaker episodic hydrodynamic
regimes. The bones showed little evidence of
tooth marks, cuts or punctures that indicated
damage by mammalian carnivores. Low relative
abundances of carnivore species further
supported a conclusion that the Grant Hall fossil
assemblage was not a large predator
accumulation.
Moriarty et al. (2000) cautioned that the
bones and sediments in Grant Hall might have
been reworked from upslope. This study found
that the fossils were well preserved and there
was a lack of evidence for substantial transport
and reworking of bones and sediments; thus, it
was concluded that the fossils were
contemporaneous with the dates assigned to the
encapsulating speleothems.
Nature of the assemblage: palaeoecology and
palaeoenvironmental reconstruction.
The Grant Hall fossil assemblage contains a
diverse large mammal fauna dominated by the
subfamilies Macropodinae and Sthenurinae. The
high frequency of macropodids is a common
feature of other fossil assemblages at the
Naracoorte Caves (Wells et al. 1984, Brown &
Wells 2000). Their frequent entrapment is partially
a function of their bounding locomotion, an
inability to move their hind limbs separately and
their comparatively small weaker forelimbs which
together make it more likely that they fall in and
less likely they can climb out. The other species,
such as wombats and koalas are perhaps less
susceptible to falling unaware down holes and
have greater mobility to manoeuvre themselves
out of partial entrapment at the entrance.
However, the substantially greater relative
abundance of macropodids in comparison to
other groups, did suggest they made up a large
proportion of the overall herbivore palaeo-
community.
Approximately 65% of the large mammalian
herbivores were browsers. For example, Wallabia
bicolour, the most abundant kangaroo in the
deposit, feeds mainly on tree leaves and soft
browse and inhabits the wetter forested
environments of eastern Australia (Strahan 2000).
The greatest macropodid diversity occurred
within the extinct Sthenurinae; and they have
been considered browsers based on the nature
of their cranial and dental morphologies (Prideaux
1999, 2004). Their low crowned bilophodont
molars and well-developed jaw musculature,
suggests these large ‘bulky’ and ‘robust’
kangaroos fed on a variety of tree leaves and
tougher shrubby browse (Prideaux 1999, 2004).
Although Macropus rufogriseous, second most
abundant kangaroo in the deposit, is a grazer
(Strahan 2000), which indicated that open patches
of grass were also present in the
palaeoenvironment. Overall, the greater relative
abundances of browsing species than grazing
species provided significant evidence for the
presence of a well-forested environment near
Grant Hall cave.
Small ground dwelling and arboreal
mammals that depend on adequate undergrowth
and tree cover, such as Potorous tridactylus
(Kerr, 1792), Pseudocherius peregrinus
(Boddaert, 1795), Petaurus breviceps
(Waterhouse, 1839) and Cercartetus nanus
(Desmarest, 1818), were additional evidence for
woodland with a dense understorey close to the
cave’s entrance. The presence of rodent species
that nowadays inhabit drier environments, such
as Pseudomys apodemodies (Finlayson, 1932),
were most likely collected via avian predators,
such as owls. Tyto alba (Scopoli, 1769), a
suggested accumulator, may hunt over a 20 km
distance (M.McDowell, pers. com.) and return
prey from different microhabitats in the region.
This is a plausible explanation for why the fossil
assemblage contained species with mutually
exclusive habitat preferences.
Accompanying palaeoclimatic and
palaeobotanical evidence
The ESR date of 125 Ka (Grünet al. 2001) placed
the Grant Hall fauna within the Last Interglacial
period, and marine oxygen isotope stage 5. The
oxygen isotope (ä18O) signals from Pacific marine
cores (Williams et al. 1993, Belperio et al. 1995,
Lambeck & Chappell 2001) indicate that
interglacial environments corresponded to
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ALCHERINGA
158 FRASER AND WELLS
higher sea levels and warmer temperatures than
glacial periods do (Bowler 1976, Croke et al.
1999). If this broad model of past climatic
conditions prevailed in the Naracoorte region
during this period, then higher water tables,
resulting from higher sea levels, might have fed
the low-lying swamps in the inter dune areas.
Therefore, a mosaic of habitats could have existed
between the higher more exposed drier dunes
and the wetter low-lying swamps; which in turn
supported a diverse fauna (comprising both
‘wetter’ and ‘drier’ adapted species) within a
relatively small geographic area.
The climate reconstruction of the last 500 Ka,
based on ä18O values from speleothems (Ayliffe
et al. 1998), currently provides the most fine-
grained hydrological record of the Naracoorte
region. The ä18O record has a detailed chrono-
logical resolution based on uranium-series dating,
and it covers a longer time period than that
estimated for the Grant Hall fossil deposit. Their
climate reconstruction refers to periods of
‘effective precipitation’, which are modelled on
the complex interactions between the infiltration
of surface precipitation, evaporation and
temperature to determine net hydrological balance
and subsequent speleothem deposition in each
cave. Ayliffe et al. (1998) state that stadials and
cool interstadials were periods of the greatest
effective precipitation, whereas, interglacials,
warm interstadials and glacial maximas were
comparatively dry. If the 125 Ka age is correct,
these findings place the Grant Hall fauna in a
period of warmer and drier climatic conditions,
which is somewhat at variance with the previous
interpretation of a wetter more densely forested
palaeoenvironment based on the faunal species
analysis. Two things remain unresolved before
this can be considered a major problem. Firstly,
the 125 Ka age, based on the ESR dating of one
tooth sample (depth of ~60cm), may not
necessarily represent the age of the entire Grant
Hall deposit. Although the uranium series dates
on the encapsulating speleothems (discussed
above) confine the fossils to between 206 and 76
Ka, deposition may have occurred during a range
of climatic periods throughout those 130,000
years. We do not know exactly when the deposit
began to accumulate after 206 Ka, or what hiatus
if any existed between the top of the sediment
and the 76 Ka date. Assuming correct
stratigraphic superposition, the fossils occurring
in the top 60 cm could be younger than 125 Ka,
and may have been deposited during the wetter
periods that Ayliffe et al. (1998) state occurred at
around 90-95 Ka and 105-115 Ka. Also assuming
the 76 Ka flowstone marks the cessation of
sedimentation, then the top 60 cm represents
about 50,000 years; yet, the two pale sandier sub-
layers (at depths of approximately 20 cm and 50
cm) might represent hiatuses in sedimentation
during this time. Secondly, it is difficult to
corroborate the interpretations arising from two
lines of evidence that have different levels of
resolution. In particular, the Grant Hall faunal
assemblage has ‘low resolution’, being a
collection of bones that may have accumulated
over long periods (perhaps even sporadically),
from animals living nearby or in the wider
surrounding region. In contrast, the speleothem
records have a precise and detailed chronology,
and represent a single high resolution climate
record for one specific cave location. Both
provide sound evidence for palaeoecological
reconstruction of Grant Hall. However, until more
fossils from subsequent levels are dated, it may
be too soon to conclude that palaeoecological
interpretations based on faunal analysis are
entirely inconsistent with the high resolution
palaeoclimate records from the cave speleothems.
Botanical remains have not been preserved
in the cave sediments at Naracoorte, and
currently, there is a lack of palaeobotanical
evidence from the immediate vicinity of the caves
and the wider Naracoorte area. This paucity in
data limits the palaeoecological reconstructions
that can be made for this time period. Pollen from
Lake Wangoom (Harle et al. 1999) and near-shore
marine core E55-6 (Harle 1997), however, provide
the closest regional palaeobotanical evidence
pertaining to the time of bone deposition in Grant
Hall; respectively, these sites are approximately
220 and 180 km from the Naracoorte caves. The
marine E55-6 core was dated by U/Th methods
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ALCHERINGA 159LATE PLEISTOCENE OF GRANT CAVE, NARACOORTE
and dates for parts of the Lake Wangoom core
that were past the radiocarbon limit (~45 –50 ka)
were extrapolated from pollen comparisons with
core E55-6. Harle et al. (1997, 1999) observed
that the marine oxygen isotope stage 5e
corresponded to peaks in Eucalyptus pollen and
high proportions of rainforest and fern taxa, which
suggested the presence of wet sclerophyll forest
and some rainforest within the region. In contrast,
the cooler and perhaps drier glacial periods had
greater proportions of woody herbaceous and
grass taxa. Although these pollen cores were not
‘Naracoorte specific’, they have provided a
general overview of regional vegetation during
this period and contributed to the reconstruction
of the Grant Hall palaeoenvironment.
Despite the paucity in palaeobotanical
evidence, and the difficulties in placing the age
of the entire Grant Hall fauna within a more precise
palaeoclimatic setting, it is still reasonable to
claim that during the bone accumulation period
the environment around the cave supported a
densely wooded forest with small more-open
patches of shrubs and thick grasses, a productive
soil horizon, and a diverse faunal community of
predominantly browsing species.
Comparison between Cathedral Cave and
Fossil Chamber in Victoria Fossil Cave,
Naracoorte
Uranium-series dating of speleothems associated
with cave sediments have established the
chronologies of fossil deposits at Naracoorte.
The age of the Cathedral Cave deposit was
confined between 160 Ka and 279 Ka, and Fossil
Chamber in Victoria Fossil Cave (VFC) was
assigned a minimum age of 213 Ka by Moriarty
et al. (2000). The age of the Grant Hall deposit
was confined between 206 and 76 Ka (Ayliffe &
Veeh 1988), and ESR dating of teeth from the
lower layers provided an average age of 125 Ka
(Grün et al. 2001). The Grant Hall fossil deposit
is therefore one of the youngest late Pleistocene
cave sites to be investigated within the
Naracoorte area, and it is unlikely that its age
overlaps with the other deposits.
The dark red/brown sandy clay sediments
in Grant Hall contrast with the sandier pale
sediments of Cathedral Cave and Fossil Chamber.
The Pliocene or Pleistocene Parilla sands were
identified as a possible source for the three
sediment layers found in Cathedral Cave (Brown
& Wells 2000). Moriarty et al. (2000) concluded
that the Grant Hall sediments were derived from
a terra-rossa palaeosol, possibly indicative of a
wet climate. However, a more detailed analysis
of the Grant Hall sediments would be needed to
identify specific source horizons outside the
cave. Different sediments might be indicative of
the environmental conditions prevailing at time
of deposition.
Pitfall trapping has been suggested as the
main mechanism responsible for bone
accumulation in many of the fossil deposits at
Naracoorte. VFC Fossil Chamber and Cathedral
Cave deposits have trapped faunas for longer
time periods than Grant Hall, including both
glacial and interglacial periods. All fossil
assemblages are dominated by Macropodine and
sthenurine kangaroos, with smaller represent-
ations of other large mammal taxa, for example:
wombats, Thylacinus cynocephalus,Thylacoleo
carnifex and Zygomaturus trilobus (Wells et al.
1984, Brown & Wells 2000, Reed & Bourne 2000).
In respect to the Macropodine faunas, Grant Hall
has two unique features that distinguish it from
the other deposits. The first is the absence of
the large grazing kangaroos Macropus
giganteus and M. fuliginosus, which are
abundant in the Fossil Chamber deposits of VFC
and Cathedral Cave. The second is the relative
abundances of the small macropodines
Macropus rufogriseous and Wallabia bicolour.
In Grant Hall they are nearly identical; 17.4 and
18.3% respectively. In VFC Fossil Chamber the
ratio is 200:6 (Wells et al. 1984) and in Cathedral
Cave they make up only 7 and 1%, respectively,
of the large mammal group.
Summary
The fossil deposit in Grant Hall contains a diverse
fossil assemblage suggestive of a wetter and
more densely vegetated palaeo-environment
within the Last Interglacial period, than currently
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ALCHERINGA
160 FRASER AND WELLS
exists in the region today. The deposit is well
constrained in time, to an age of approximately
125 Ka; a time period not previously investigated
at the Naracoorte Caves. Hence this investigation
was partially motivated to amend the paucity of
palaeontological data for this time period.
Although the overall species composition
of the Naracoorte faunas appears to have
changed little throughout the Middle to Late
Pleistocene (Moriarty et al. 2000), differences in
the relative proportions of the Macropodidae
species might be one of the more significant
palaeoenvironmental indicators in these
deposits. In this respect, the Grant Hall fauna
contains greater relative abundances of
browsing kangaroos than the other Naracoorte
cave faunas. This feature sets Grant Hall apart.
This project provides evidence that at the
peak of the Last Interglacial the Naracoorte
region supported a diverse suite of large
browsing herbivores and their predators, which
is noticeably different to the faunal communities
of today. Extinct megafauna species, such as
the Sthenurine kangaroos, Zygomaturus trilobus
and Thylacoleo carnifex, which ‘characterise’
many Pleistocene fossil assemblages Australia-
wide (Murray 1991) were present at Naracoorte.
Moreover, this study reveals that at 125 Ka these
species still formed a diverse and dominant
component of the mammalian faunal community.
The chronological controls provided by
both uranium series and electron spin resonance
dating, together with a rigorous taphonomic
investigation has enabled a much more detailed
analysis and interpretation of this important
deposit. In conclusion, the faunal analysis of
the Grant Hall fossil assemblage provided
evidence that faunas within the Naracoorte
Caves region fluctuated in both species
composition and relative abundances in
response to changing climatic and environmental
regimes throughout the mid to late Pleistocene.
Further investigation into younger cave
deposits, utilising absolute dating methods and
taphonomic analyses, are needed to constrain
the chronology of large mammal faunal
extinctions and to trace the evolution and
structure of the mammalian community in this
southern region.
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