Quaternary Science Reviews 37 (2012) 38e47

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Quaternary Science Reviews

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Man and megafauna in Tasmania: closing the gap

Richard Gillespie a, b, *, Aaron B. Camens c, Trevor H. Worthy d, Nicolas J. Rawlence e, f, Craig Reid g,
Fiona Bertuch h, Vladimir Levchenko h, Alan Cooper e
a
Centre for Archaeological Science, School of Earth and Environmental Sciences, University of Wollongong, Wollongong, NSW 2522, Australia
b
Division of Archaeology and Natural History, School of Culture, History and Language, Australian National University, Canberra, ACT 0200, Australia
c
School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia 5005, Australia
d
Department of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW 2052, Australia
e
Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia 5005, Australia
f
Department of Earth and Ocean Sciences, University of Waikato, Private Bag 3105, Hamilton 3240, New Zealand
g
Queen Victoria Museum and Art Gallery, PO Box 403, Launceston, Tasmania 7250, Australia
h
Institute for Environmental Science, Australian Nuclear Science and Technology Organisation, Locked Bag 2001, Kirrawee DC, NSW 2232, Australia

a r t i c l e i n f o a b s t r a c t

Article history: Recent discussion on the late Pleistocene extinction of the Australian megafauna has revolved around
Received 9 September 2011 interpretation of several key fossil sites in Tasmania. It has been suggested that humans did not arrive in
Received in revised form Tasmania until after the megafauna became extinct, or did not hunt now extinct megafauna, and
27 November 2011
therefore that humans cannot be implicated in the extinctions. Radiocarbon results from these sites
Accepted 11 January 2012
Available online 10 February 2012
indicate that the youngest extinct megafauna are close to charcoal ages from the oldest archaeological
deposits, although difficulties have arisen in establishing chronologies because most relevant sites have
ages near the limit for radiocarbon analysis.
Keywords:
Megafauna
We report a series of new radiocarbon ages, d13C, d15N and C:N ratios on collagen and dentine fractions
Radiocarbon from skeletal remains in the Mount Cripps karst area and the Mowbray Swamp, both in northwestern
Collagen Tasmania, and discuss the reliability of ages from these and other sites. We also report the discovery of an
Extinction articulated Simosthenurus occidentalis skeleton at Mt Cripps, that represents only the second directly-
Australia dated extinct megafaunal taxon with a reliable age <50 ka cal BP from Tasmania.
Our results suggest that C:N ratios measured on collagen or dentine are not an infallible guide to
radiocarbon age reliability. We confirm previous reports of a temporal overlap between the megafaunal
and archaeological records in Tasmania, but the presence of archaeological evidence and megafauna with
the same age at the same site has not yet been demonstrated. At least two megafaunal taxadthe now-
extinct Protemnodon anak and a giant Pleistocene form of the extant Macropus giganteusdwere still
present in Tasmania after 43 ka, when human crossing of the Bassian landbridge from mainland Australia
first became sustainable.
Ó 2012 Elsevier Ltd. All rights reserved.

1. Introduction mainland Australia’s large-bodied Pleistocene fauna became

Early human occupation of mainland Australia in the 50e40 ka
age range is well-documented, often with multi-method dating;
notable examples include Lake Mungo in the southeast (Bowler
et al., 2003), Devil’s Lair in the southwest (Turney et al., 2001),
and in the northwest at Carpenter’s Gap (O’Connor, 1995) and Riwi
(Balme, 2000). Around this same 50e40 ka interval, the majority of
* Corresponding author. Centre for Archaeological Science, School of Earth and
Environmental Sciences, University of Wollongong, Wollongong, NSW 2522, Aus-
tralia. Tel.: þ61 2 6677 9500.
E-mail address: dizzy@better.net.au (R. Gillespie).
extinct (e.g. Roberts et al., 2001; Miller et al., 2005; Grün et al.,
2008; Prideaux et al., 2010). This temporal coincidence has stimu-
lated hotly-debated, if largely inconclusive, argument concerning
the sequence, timing and causes of the Australian megafaunal
extinctions (e.g. Gillespie et al., 2006; Brook et al., 2007; Field et al.,
2008).
Several aspects of the Australian debate are similar to those
concerning the North American extinctions (e.g. Grayson and
Meltzer, 2003; Fiedel and Haynes, 2004), with one significant
difference: among the large number of sites bridging the Rancho-
labrean Termination (Haynes, 2008), there are at least 14 well-
dated “elephant kill sites” in the North American record where
both directly-dated extinct taxa and associated subsistence

0277-3791/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.quascirev.2012.01.013
 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47
archaeology of the same age occur in the same stratigraphic layers
(e.g. Grayson and Meltzer, 2002; Surovell and Waguespack, 2008).
Australia has no known sites with these attributes, and conse-
quently the debate is reduced to polarised views based on the
contents of a very small number of either fossil or archaeological
sites in the appropriate age range. The island state of Tasmania,
periodically connected to mainland Australia during sea level
lowstands, potentially offers an interesting test for late Pleistocene
extinction hypotheses.
Lambeck and Chappell (2001) determined that the Bass Strait
between mainland Australia and Tasmania was bridged w43 ka,
allowing pedestrian human crossing for the first time and
suggesting a significantly shorter overlap period for humans and
megafauna in Tasmania than on mainland Australia. This accords
well with the oldest Tasmanian archaeological sites so far docu-
mented, Warreen Cave (Allen, 1996) and Parmerpar Meethaner
(Cosgrove, 1995), which have maximum calibrated radiocarbon
(14C) ages measured on charcoal ca. 40e38 ka cal BP. Compared
with mainland Australia, the extinct megafauna record from Tas-
mania is depauperate, probably including only Zygomaturus trilo-
bus, Palorchestes azael, Metasthenurus newtonae, Simosthenurus
occidentalis, Protemnodon anak, Thylacoleo carnifex and Mega-
libgwilia sp. (Turney et al., 2008) plus a giant form of the extant
Macropus giganteus.
With the first defendable direct 14C age measurements on
extinct Australian marsupial bones, ca. 43e41 ka cal BP on
Protemnodon from Mt Cripps, Turney et al. (2008) suggested that
Tasmania may have been the last refuge for the Australian
Melbourne
Bass Strait
Mowbray
Swamp
Mt Cripps
Parmerpar
Meethaner
ORS 7
Pallawa Trounta
Warreen Cave
Nunamira
Bone Cave
144°E 146°E
Fig. 1. Map showing the location of Tasmanian fossil sites (red filled circles) and archaeological sites (white filled circles) discussed in the text, with an approximate contour for the
Bassian landbridge w43 ka (modified from Cosgrove et al., 2010; Lambeck and Chappell, 2001). (For interpretation of the references to colour in this figure legend, the reader is
referred to the web version of this article.)
39
megafauna. But despite the analysis of many thousands of bone
fragments, no skeletal remains of any extinct taxon have been
found in Tasmanian archaeological sites (Cosgrove et al., 2010), an
observation that we discuss below. Reliable dating of Tasmanian
archaeological and palaeontological sites has been limited due to
these sites being at or near the limit of radiocarbon dating, adding
to the difficulties in establishing the provenance of sediments used
for OSL dating of key sites (Cosgrove et al., 2010).
For this study, we measured 14C ages, stable C and N isotopes,
and C:N ratios on collagen and dentine fractions isolated from the
skeletal remains of three extant and four extinct marsupial taxa
recovered from the Mt Cripps and Mowbray Swamp fossil sites in
northwestern Tasmania. We compare our directly-dated marsupial
ages with previous results from Turney et al. (2008) and Cosgrove
et al. (2010), assessing their 14C age reliability and any temporal
overlap with the existing Tasmanian archaeological record. The
locations of fossil and archaeological sites in Tasmania discussed
here are shown in the Fig. 1 map.
2. Materials and methods
Table 1 shows details of bones and teeth sampled for this study
from Mt Cripps and Mowbray Swamp, which are housed at the
Queen Victoria Museum and Art Gallery (QVMAG) in Launceston
and the Australian Centre for Ancient DNA (ACAD) in Adelaide.
Collagen content of the bones was estimated using a ninhydrin
method on decalcified and hydrolysed bone samples, compared
with similarly-prepared aliquots from a modern Wallabia bicolor
38°S
Landbridge
~43 ka
40°S
Launceston
42°S
JF154, JF155,
Ultimate Cave,
Titan’s Shelter
Hobart
0 100 km
148°E
40 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47

Table 1 Lastly, all fractions were freeze dried before combustion, graphi-
Bones and teeth analysed in this study, showing sample location, museum accession tisation and AMS radiocarbon analysis using standard methods at
numbers, body part and sample ID used in subsequent tables; taxa marked with an
asterisk are extinct megafauna.
ANSTO (Fink et al., 2004) or the ORAU (Bronk Ramsey et al., 2004).
Separate sub-samples of each fraction were combusted in a coupled
Location Species Body part QVM No. Sample ID elemental analyser-isotope ratio mass spectrometer system for
Mt Cripps Simosthenurus upper I1 frag. 2009 GFV:0002 MCSt-1 measurement of d13C, d15N and atomic C:N ratios.
(CP222) occidentalis*
We used the 2s sample significance and pooled mean age tools
Mt Cripps Protemnodon anak* L tibia 2001 GFV:2 MCP-1
(CP213) in CALIB 6.0 (Stuiver et al., 2010) to compare conventional 14C ages
L femur 2001 GFV:40 MCP-2 from the different fractions prepared in this study. Those ages
L lower I1 2011:GFV:1 MCP-3 considered acceptable were calibrated with OxCal 4.1 (Bronk
Thylogale billardierii L tibia 2001 GFV:68 MCT-1 Ramsay, 2009) using the IntCal09 atmospheric data in Reimer
Vombatus ursinus R femur 2001 GFV:30 MCV-1
upper P3 2011:GFV:2 MCV-2
et al. (2009).
Sarcophilus harrisii R femur 2001 GFV:39 MCS-1
Mowbray Zygomaturus trilobus* rib frag. 1992 GFV:148 MSZ-1 3. Results and discussion
Swamp
femur frag. 1992 GFV:181 MSZ-2
Conventional radiocarbon ages, d13C, d15N and C:N ratios
Palorchestes azael* femur frag. 2007 GFV:22 MSPal-1
measured on our Tasmanian marsupial samples, and on the
radiocarbon bone standard VIRI sample E (a mammoth bone from
tibia (Gillespie and Brook, 2006). Examined under 10 binocular Quartz Creek, Yukon; Scott et al., 2009) processed with our samples
magnification, the decalcified bone samples from Mt Cripps and at ANSTO, are presented in Table 2. We note that C:N ratios for
Mowbray Swamp exhibited acceptable collagen pseudomorphs collagen and gelatin fractions from these bones and teeth are all
(Stafford et al., 1987), with collagen content ranging from 3 to 17% very similar, and all are within the acceptable range of 2.9e3.5
by weight of original bone; these were analysed at the Australian (DeNiro, 1985; Brock et al., 2007).
Nuclear Science and Technology Organisation (ANSTO) in Sydney.
The teeth, which yielded 1.9e4% dentine, were analysed at the 3.1. Standard bone
Oxford Radiocarbon Accelerator Unit (ORAU), UK. Bones and teeth
were processed using methods described in Stafford et al. (1991) Radiocarbon ages for our three independent preparations of
and Brock et al. (2007) to prepare a subset of the following ultrafiltered gelatin from the VIRI E standard bone are statistically
fractions: identical. Their pooled mean age of 39,430  310 BP is not signif-
icantly different from the VIRI consensus age of 39,305  121 BP,
rawC e cold decalcification: 4  C for several days, daily acid suggesting that the methods employed here produce results
change consistent with best practice for late Pleistocene bone.
coldFG e cold decalcification, gelatinisation, 0.45 m PTFE
filtration 3.2. Mt Cripps
stdUG e Oxford protocol: room temp. ABA, gelatinisation, Eezi-
filter, Vivaspin-15 ultrafilter Collagen and gelatin fractions from Mt Cripps bones are pale
coldUG e same except cold decalcification of bone fragments straw colour, similar to those from the well-preserved standard

Table 2
Analytical results for Mt Cripps and Mowbray Swamp skeletal remains, and the VIRI E standard bone, showing chemical fraction (see text for details), percent Modern C,
conventional 14C age, stable C and N isotope ratios, and atomic C:N ratio; taxa marked with an asterisk are extinct megafauna. Data from the AMS facilities at ANSTO, Sydney,
and Oxford, UK; n/a ¼ not available.

Site Taxon Sample ID Fraction Lab No. % Mod. C Conv. 14C age d13C d15N Atomic C:N
measured (1s BP) (0.4&) (0.4&) ratio (0.2)
Mt Cripps CP222 Simosthenurus MCSt-1 stdUG OxA-17143 0.39  0.05 44,500  1000 22.9 n/a 3.2
occidentalis *
Mt Cripps CP213 Protemnodon anak * MCP-1 stdUG OZM081 1.08  0.04 36,350  300 24.3 2.7 3.4
MCP-1/2 coldUG OZM739 1.00  0.05 37,010  410 24.0 2.7 3.4
Protemnodon anak * MCP-2 stdUG OZM082 0.67  0.03 40,170  370 24.2 2.6 3.3
MCP-2/2 coldUG OZM744 0.72  0.06 39,660  680 23.8 2.7 3.3
MCP-2/4A rawC OZN266U1 0.92  0.04 37,670  360 23.9 2.9 3.3
MCP-2/4B coldFG OZN266U2 0.83  0.03 38,520  300 23.6 2.5 3.3
Protemnodon anak * MCP-3 stdUG OxA-22701 0.59  0.10 41,200  1400 22.1 2.5 3.3
Thylogale billardierii MCT-1 stdUG OZM083 0.99  0.04 37,080  330 24.0 2.3 3.5
Vombatus ursinus MCV-1 stdUG OZM736 0.70  0.04 39,900  470 23.9 3.8 3.3
Vombatus ursinus MCV-2 stdUG OxA-22702 0.53  0.10 42,000  1600 22.2 3.7 3.1
Sarcophilus harrisii MCS-1 stdUG OZM084 0.65  0.03 40,510  380 22.6 2.0 3.3

Mowbray Swamp Zygomaturus trilobus * MSZ-1 stdUG OZM085 1.54  0.04 33,510  210 21.9 5.0 3.3
MSZ-1/2 coldUG OZM740 1.05  0.06 36,570  460 21.7 5.0 3.3
MSZ-1/4 coldFG OZN265 1.38  0.05 34,410  300 21.5 4.4 3.3
Zygomaturus trilobus * MSZ-2 coldUG OZM737 0.29  0.02 46,810  560 20.6 6.2 3.3
MSZ-2/4 coldFG OZN264 0.50  0.03 42,500  480 20.4 6.3 3.4
Palorchestes azael * MSPal-1 coldUG OZM738 1.05  0.04 36,580  310 22.3 3.2 3.4
MSPal-1/2 coldFG OZN263 1.14  0.04 35,920  290 22.3 3.2 3.3

Quartz Ck., Yukon Mammuthus sp. * VIRI E-1 stdUG OZI812U1 0.75  0.07 39,350  740 20.4 n/a n/a
(Reference Standard) VIRI E-2 stdUG OZI812U2 0.72  0.04 39,620  450 20.8 n/a n/a
VIRI E-3 stdUG OZI812U3 0.76  0.05 39,220  530 20.7 n/a n/a
 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47
bone. We compared four fractions prepared from Protemnodon
bone MCP-2, finding that the ages on standard and cold-decalcified
ultrafiltered gelatin were statistically identical (pooled mean age
40,050  325 BP), while the ages on raw collagen and 0.45 m PTFE
filtered gelatin fractions from the same bone were significantly
younger and consequently rejected. The juvenile Protemnodon tibia
(MCP-1, see photo in Fig. 2A) also produced identical ages for
standard and cold-decalcified ultrafiltered gelatin, with pooled
mean age 36,580  240 BP. We accept ages on all standard or cold-
decalcified ultrafiltered gelatin fractions from samples MCP-1,
MCP-2, MCP-3, MCT-1, MCS-1, MCV-1 and MCV-2 (cave CP213),
and MCSt-1 (cave CP222).
Measurements on Protemnodon femur MCP-2 and the isolated
lower incisor MCP-3 are identical, and their pooled mean age of
40,110  320 BP is significantly older than the MCP-1 tibia mean
age. Ages on bone MCT-1 from the extant taxon Thylogale has the
same age as Protemnodon bone MCP-1, and samples from the extant
taxa Sarcophilus (MCS-1) and Vombatus (MCV-1, MCV-2) have the
same age as Protemnodon bone MCP-2. Thus we have shown that
both extant and extinct species from Mt Cripps CP213 have a similar
age range ca. 36e42,000 BP.
Fig. 2. (A) Protemnodon anak juvenile tibia MCP-1 (QVM:2001 GFV: 2), scale bar equals
100 mm; (B) Simosthenurus occidentalis adult skull MCSt-1 (QVM:2006 GFV: 002),
scale bar equals 100 mm; (C) decalcified collagen from Zygomaturus trilobus rib MSZ-1
(QVM:1992 GFV:148), note internal browneblack filaments of possible organic
contamination, scale bar equals 1 mm.
41
Our oldest acceptable radiocarbon age of 44,500  1000 BP was
measured on an upper incisor fragment (MCSt-1) of an adult S.
occidentalis skull from cave CP222 (see photo in Fig. 2B), part of an
articulated skeleton to be described in detail elsewhere (Camens
and colleagues, manuscript in prep.), and the second directly-dated
extinct marsupial species from Mt Cripps.
3.3. Mowbray swamp
Given previous unsuccessful attempts to find collagen in
megafaunal bones and teeth from Lancefield Swamp and several
other mainland Australian fossil sites (Gillespie et al., 1978;
Gillespie and Brook, 2006; Gillespie, unpubl. data), the presence of
collagen in extinct marsupial bones from Mowbray Swamp was
unexpected; however, we found 2e4% collagen in samples MSZ-1,
MSZ-2 and MSPal-1. These Z. trilobus and P. azael bones were
described by Gill and Banks (1956) as “.various shades of
brownish grey and grey, being stained presumably by the peat and
probably such iron as is presently being chemically reduced by the
decomposing vegetable matter.” Viewed under 50 magnification,
thin brown-black filaments are visible within the translucent
cold-decalcified collagen pseudomorph from rib bone MSZ-1 (see
photo in Fig. 2C), and the gelatin fractions we prepared from these
bones are also darkly stained.
Radiocarbon ages on the gelatin preparations are scattered. Our
comparison of three fractions from bone MSZ-1 found all ages to be
significantly different, with the oldest age of 36,570  460 BP on
cold-decalcified ultrafiltered gelatin. Cold-decalcified 0.45 m PTFE
filtered gelatin yielded a younger age than ultrafiltered gelatin for
bone MSZ-2, whereas these fractions from MSPal-1 have statisti-
cally identical ages (pooled mean age 36,230  210 BP).
Ages on wood and peat from the Mowbray Swamp (Gill and
Banks, 1956; van de Geer et al., 1986) exhibit the same asymp-
totic pattern as those from the nearby Pulbeena Swamp (Colhoun
et al., 1982). As discussed by Chappell et al. (1996), a radiocarbon
‘event horizon’ exists at the limits of available decontamination and
measurement technology, and when a set of 14C dates cluster near
an asymptote, as they do for Mowbray Swamp, there is a high
probability that the ages are unreliable. We interpret these wood
and peat 14C results to mean that the sediments overlying the layers
containing the extinct marsupials sampled here are >52,000 BP
and could be much older. This does not necessarily imply that the
bones are the same age as the peat layers in which they were found,
nor that the three bones necessarily have the same true age. These
low-lying peat swamps were drained by farmers early in the 20th
century, and when sampled by van de Geer et al. (1986) the
thickness of peat at Mowbray Swamp had decreased from more
than 4 m to w2 m. While the expectation would be that humic and
fulvic acids from the peat would contribute older carbon, the
contamination we observe appears to be young.
Even though C:N ratios of gelatin fractions from these Zygo-
maturus and Palorchestes bones are all within the acceptable range,
regardless of chemistry variations, we reject all our Mowbray
Swamp results. The bones contain browneblack organic contami-
nation that so far has proven impossible to completely remove
using the usual acid and alkali solutions, several combinations of
organic solvents, or the Oxford ultrafiltration protocol. Even our
oldest result of 46,810  560 BP on bone MSZ-2 may seriously
underestimate the true age.
3.4. Stable C and N isotopes
The d13C and d15N ratios measured on bone collagen/gelatin
fractions prepared in this study (see Table 2 and Fig. 3) exhibit
several interesting features:
42 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47
A B
Zygomaturus trilobus
Palorchestes azael
Protemnodon anak
Simosthenurus occidentalis
Thylogale billardierii dentine
Vombatus ursinus
Sarcophilus harrisii
M. rufogriseus
M. giganteus (large morph)
Simosthenurus sp.
-25 -24 -23 -22 -21
δ13C (‰)
Fig. 3. (A) d13C (0.4& PDB) values for bone gelatin or tooth dentine on taxa with ages >35 ka cal BP, diagonal dashed line separates northern (Mt Cripps, Mowbray Swamp) from
southern sites (Titan’s Shelter, JF154, JF155, Ultimate Cave, Warreen Cave). Rectangles are Mt Cripps samples (the carnivore Sarcophilus harrisii has star shape), circles are Mowbray
Swamp samples, triangles are from the southern sites; red filled shapes are extinct taxa, dark green filled shapes are extant taxa, light green shapes are M. giganteus (large morph).
(B) Plot of available d13C v. d15N (0.4& AIR) values for bone gelatin from herbivorous taxa with ages >40 ka cal BP; symbols as above. (For interpretation of the references to colour
in this figure legend, the reader is referred to the web version of this article.)
(1) the small amount of organic carbon contamination that
reduces the age of contaminated bone gelatin appears not to
significantly affect the stable isotope or C:N ratios;
(2) the observed difference found between bone from the only
carnivorous marsupial sampled here (Sarcophilus harrisii,
d13C ¼ 22.6&) and the similarly-aged herbivorous marsupial
bones from Mt Cripps (d13C range 23.8 to 24.2&) is as ex-
pected given their trophic level;
(3) where measurements are available on bones and teeth from
the same taxon, e.g. Protemnodon samples MCP-3 (tooth) and
MCP-2 (bone) and Vombatus MCV-2 (tooth) and MCV-1 (bone),
dentine gelatin has less negative d13C ratios than bone gelatin;
(4) the stable isotope signatures of our herbivore bones, regardless
of chemistry variations, cluster in two distinct geographic
groups; a third cluster is apparent when previous results from
the southern fossil sites Titan’s Shelter, JF154, JF155, Ultimate
Cave (Cosgrove et al., 2010) are included, shown in Fig. 3B.
The stable isotope variations in these Tasmanian herbivores are
likely correlated with latitude, regional climate, water availability,
elevation and temperature (e.g. DeNiro and Epstein, 1978;
Bocherens et al., 1995; Richards et al., 2002; Murphy et al., 2007).
Marsupial lifestyle may also be important, for example the dietary
changes reflected in d13C values of bone collagen from pouch young
to adult kangaroos (Witt and Ayliffe, 2001). Adult body mass range
for the herbivores sampled here (e.g. Murray, 1991; Helgen et al.,
2006; van Dyck and Strahan, 2008) extends from the very large
500e700 kg Z. trilobus and w300 kg P. azael at the coastal lowland
Mowbray Swamp (10e15 m above sea level, van de Geer et al.,
1986), through the 97e136 kg S. occidentalis, 122e153 kg P. anak
and the small 4e7 kg Thylogale billardierii at the upland Mt Cripps
karst (355 masl, Turney et al., 2008). Our observation that samples
from sites cluster together regardless of taxa suggests that site-
specific characteristics affect the stable isotope composition more
than intrinsic species-specific factors. Exploring the relationships
between the stable isotope composition of marsupial skeletal
remains with body mass, vegetation and climatic variables is
beyond the scope of this paper and will not be discussed further.
3.5. Summary of results
We report acceptable radiocarbon ages on two extinct and three
extant marsupial taxa from Mt. Cripps: five ages on two bones and
one tooth of the extinct P. anak; one age on a tooth of the extinct S.
7
Mowbray Swamp
6
5
Mt Cripps
δ15N (‰)
4
3
2
1 Titan’s Shelter, JF155,
JF154, Ultimate Cave
0
-20 -25 -24 -23 -22 -21 -20
δ13C (‰)
occidentalis; one age on bone and one age on a tooth of the extant
Vombatus ursinus; and one age each on bones of the extant T. bill-
ardierii and S. harrisii. These new 14C ages, which cluster in the 2s
calibrated range 40.9e49.8 ka cal BP, are listed in Table 3 and
graphed as probability plots with median age in Fig. 4.
3.6. Comparison with previous Tasmanian marsupial results
Turney et al. (2008) reported inter-laboratory comparisons on
two Protemnodon bones from different individuals found in cave
CP213 at Mt Cripps. They accepted the ages on bones processed
using the standard ultrafiltration protocol at the ORAU, and rejected
results on the same bones processed at the University of Wollon-
gong using a step-combusted gelatin method because they yielded
significantly younger ages. We concur, and also reject their result
on a Protemnodon bone from Titan’s Shelter processed with the
step-combusted gelatin procedure because the C:N ratio of 2.8 is
below the acceptable range (Brock et al., 2007). Our two ages on the
Protemnodon left tibia sample MCP-1 are statistically the same as
the younger Protemnodon right tibia age accepted by Turney et al.
(2008) from cave CP213. The pooled mean age for these three
measurements on one individual at 36,430  190 BP
(41.5  0.4 ka cal BP) is the youngest for any Tasmanian extinct
megafauna.
Acceptable ages on our femur sample MCP-2 and tooth MCP-3,
both Protemnodon from cave CP213, with pooled mean age
40,110  320 BP (44.1  0.6 ka cal BP), are significantly older than
our ages on MCP-1 and both tibia analysed by Turney et al. (2008),
indicating a third individual. The combined results from four taxa
suggests that the deposit spans 41e44 ka (see Table 3 and Fig. 4),
and we agree with Turney et al. (2008) that there is no conflict
between these calibrated 14C ages and the 36  3 ka OSL age on
sand from the nasal cavity of one of the Mt Cripps Protemnodon
skullsdthe OSL age has to be younger because the sand was
necessarily deposited some time after that animal died.
Cosgrove et al. (2010) reported inter-laboratory comparisons on
bones from cave sites in south central Tasmania, and we agree with
their acceptance of ca. 41e44 ka cal BP ages on M. giganteus titan
bones from Ultimate Cave and Titan’s Shelter square F, which have
C:N ratios within the guidelines (Brock et al., 2007). However,
Macropus titan Owen, 1838 inhabited a similar niche (Poole, 1982),
was synonymised with the extant taxon M. giganteus by Dawson
and Flannery (1985), and has since been treated as a synonymous
giant Pleistocene morph (e.g. Roberts et al., 2001; Easton, 2006;
 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47 43

Table 3
Radiocarbon ages on extant and extinct fauna, and archaeological charcoal from Tasmania. Conventional 14C ages were calibrated with OxCal 4.1 (Bronk Ramsay, 2009) using
the IntCal09 atmospheric dataset in Reimer et al. (2009), showing the 2s calibrated age range and median calibrated age.

Location Taxon or material dated Lab No. Conv. 14C age Cal. age range Median age Ref.
(1s BP) (2s cal BP) (ka cal BP)
Fossil Sites
Mt Cripps CP222 Simosthenurus occidentalis * OxA-17143 44,500  1000 49,840e46,000 47.8 1
Mt Cripps CP213 Protemnodon anak * OZM081 36,350  300 41,990e40,930 41.5 1
OZM739 37,010  410 42,490e41,240 41.9 1
OZM082 40,170  370 44,760e43,370 44.1 1
OZM744 39,660  680 44,750e42,680 43.7 1
OxA-22,701 41,200  1400 47,170e42,560 44.8 1
OxA-16417 36,200  300 41,900e40,740 41.3 2
OxA-16418 37,920  340 43,010e41,920 42.5 2
Vombatus ursinus OZM736 39,900  330 44,650e43,080 43.9 1
OxA-22702 42,000  1600 48,800e41,620 45.5 1
Thylogale billardierii OZM083 37,080  330 42,450e41,380 41.9 1
Sarcophilus harissii OZM084 40,510  380 45,060e43,650 44.4 1
Titan’s Shelter M. giganteus (large morph) CAMS-128558 37,650  470 42,990e41,620 42.3 3
CAMS-96368 36,210  390 41,980e40,570 41.3 3
CAMS-96369 35,610  360 41,590e39,700 40.8 3
Ultimate Cave CAMS-96367 39,530  580 44,540e42,730 43.6 3
Archaeological Sites
Warreen Cave M. rufogriseus OZF415 31,660  250 36,670e35,280 36.1 3
Vombatus ursinus OZF416 30,610  350 36,270e34,600 35.1 3
charcoal Beta-46873/ETH-8509 31,610  370 36,720e35,130 35.9 4
charcoal Beta-46872/ETH-8508 27,160  250 31,790e31,080 31.4 4
charcoal Beta-42059/ETH-8501 30,210  300 35,330e34,080 34.8 4
charcoal Beta-42122B/ETH-7665B 34,790  510 41,030e38,740 39.9 4
charcoal ANUA-11710/143B 33,170  670 39,430e36,470 37.9 3
Parmerpar Meethaner charcoal Beta-68158/CAMS-10270 33,850  450 40,090e37,280 38.7 5
charcoal Beta-68159/CAMS-10271 33,260  420 38,860e36,810 37.9 5
charcoal Beta-61608/CAMS-6050 27,780  230 32,690e31,400 32.0 5
Nunamira Cave charcoal Beta-25880 27,770  420 33,050e31,270 32.0 6
charcoal Beta-28323 28,000  720 34,430e31,160 32.4 6
charcoal Beta-25881 30,420  690 36,480e33,490 35.0 6
ORS 7 charcoal Beta-23404/ETH-3724 30,840  480 36,420e34,660 35.5 6
Bone Cave charcoal Beta-29987 29,000  520 34,680e32,080 33.6 7
charcoal Beta-37547 28,330  720 34,450e31,430 32.7 8
Pallawa Trounta charcoal Beta-44081 29,800  720 36,270e32,550 34.3 9
charcoal Beta-62285 27,250  530 32,910e30,880 31.6 9

References: 1 This work; 2 Turney et al. (2008); 3 Cosgrove et al. (2010); 4 Allen (1996); 5 Cosgrove (1995); 6 Cosgrove (1989); 7 Allen (1989); 8 Holdaway and Porch (1996); 9
Stern and Allen (1996).

Helgen et al., 2006; Webb, 2008, 2009). Given that the taxonomic
distinction of M. titan has not yet been established through rigorous
taxonomic or biomolecular analysis, we regard it as a junior
synonym of M. giganteus and not as an extinct taxon (species or
subspecies) until such an analysis is undertaken.
The two youngest M. giganteus bones analysed by Cosgrove
et al. (2010) from Titan’s Shelter square F are statistically the
same age as the youngest Protemnodon bones from Mt Cripps
analysed in this work and by Turney et al. (2008). In our chemistry
variations on Protemnodon bone MCP-2, we found that the ultra-
filtered gelatin ages were significantly older than those measured
on 0.45 m PTFE filtered gelatin and raw collagen fractions, the
opposite pattern to that observed for M. giganteus bones by
Cosgrove et al. (2010). This might be explained by improved
quality control for bone dating at the ORAU and ANSTO since the
Cosgrove samples were processed; the Oxford ultrafiltration
protocol is time-consuming, precleaning the ultrafilters can be
tricky, and degraded bones sometimes have little or no high-
molecular weight fraction (e.g. Higham et al., 2006; Brock et al.,
2007; Hüls et al., 2007).
Cosgrove et al. (2010) accept the older of two significantly
different ages on paired samples of M. giganteus bone from Titan’s
Shelter, following the standard radiocarbon judgement that the
older of two dates likely represents a more complete removal of
contamination. But they unaccountably accept the younger of two
statistically identical ages (50,600  3000 BP and
44,700  3300 BP) on a Simosthenurus sp. bone from cave JF155.
Although the C:N ratio is within the guidelines, we reject this result
because the uncertainty is greater than usual for comparable
material (such as our results) and the pooled mean age of
47,930  2220 BP is beyond the IntCal09 calibration range. All the
finite and non-finite 14C ages on bones from sites JF154 and JF155
are likely to be unreliable because they cluster on an asymptote
near the limit of the 14C method (Chappell et al., 1996); their true
age is almost certainly >50 ka cal BP.
Table 3 lists the acceptable direct 14C age determinations on the
skeletal remains of extinct and extant marsupials from Tasmanian
fossil sites, and 14C ages (mostly on charcoal) from Tasmanian
archaeological sites, showing calibrated ages in the range
30e50 ka cal BP. Fig. 4 shows the individual calibration plots and
median ages for all samples listed in Table 3, generated with OxCal
4.1 (Bronk Ramsay, 2009) using the IntCal09 atmospheric dataset in
Reimer et al. (2009).
4. Significance for Australian megafauna extinction
chronologies
Accepting multi-method chronologies supporting the first
human colonisation of greater Australia w50 ka (e.g. Brook et al.,
2007), and that a sustainable Bassian landbridge w43 ka first
gave humans pedestrian access to Tasmania (Lambeck and
Chappell, 2001), we now address the nested set of questions con-
cerning the evidence for direct interaction between humans and
megafauna.
44 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47

Archaeological Sites
Bone Cave
Beta-37547
Beta-29987
Bassian Pallawa Trounta
Landbridge Beta-62285
Beta-44081
Numamira
Beta-25880
Beta-28323
Beta-25881
ORS7
Beta-23404/ETH-3724
Parmerpar Meethaner
Beta-61608/CAMS-6050
Mt Cripps Beta-68159/CAMS-10271
Fossil Sites

OxA-17143 Beta-68158/CAMS-10270
OxA-22701 Wareen Cave
Beta-46872/ETH-8508
Extinct

OZM082
OZM744 Beta-42059/ETH-8501
OxA-16418 OZF416
OZM739 Beta-46873/ETH-8509
OZM081 OZF415
OxA-16417 ANUA-11710/143B
OZM083 Beta-42122B/ETH-7665B
Extant

OZM736
OZM084
OxA-22702
Titan’s Shelter
CAMS-96369
Extant

CAMS-96368
CAMS-128558
CAMS-96367

50 45 40 35 30
Calibrated Radiocarbon Age (ka cal BP)

Fig. 4. Diagram showing probability plots for calibrated ages of samples from Tasmanian fossil and archaeological sites, with w43 ka age for the Bassian landbridge emergence
(Lambeck and Chappell, 2001); colours for fauna as in Fig. 3, grey filled shapes are archaeological charcoal (see Table 3). Plots generated with OxCal 4.1 (Bronk Ramsay, 2009) using
the IntCal09 atmospheric dataset (Reimer et al., 2009), short black vertical bars are median probability. Graph is truncated at 50 ka cal BP by the calibration limit, and only calibrated
ages >30 ka cal BP are included; all plots have the same vertical scale, baselines were shifted to show locations.

4.1. Fossil site megafauna
The number of Australian sites with acceptable ages from
directly-dated extinct fauna is still very small compared with those
documenting the North American and Eurasian extinctions (e.g.
Haynes, 2008; Mellars and French, 2011), and issues of 14C dating
reliability for late Pleistocene megafaunal assemblages in Australia
have attracted recent attention (e.g. Miller et al., 1999; Roberts
et al., 2001; Gillespie et al., 2006; Gillespie, 2008; Turney et al.,
2008). However, unlike the North American extinction period
ca. 14e12 ka, ages in the 50e40 ka cal BP range are pushing the
limits of the 14C method, and the glaciofluvial stratigraphic record
available to constrain the age of late Pleistocene events is better
developed in Europe than it is in Australia.
Roberts et al. (2001) suggested, based on a statistical analysis of
seven sites containing articulated skeletal remains with reliable
OSL or U/Th ages <55 ka, that megafaunal extinction took place on
the Australian mainland ca. 51e40 ka. This extinction window has
been supported by several recent studies, including continent-wide
extinction of the giant flightless bird Genyornis newtoni at 50  5 ka
(Miller et al., 1999), and reports that Zygomaturus, Protemnodon,
sthenurine kangaroos and T. carnifex were among the last mega-
faunal taxa to disappear (ca. 45 ka cal BP) from the fossil record at
the Naracoorte Caves in southeastern South Australia (Pate et al.,
2002), and that Protemnodon and sthenurine kangaroos were also
the last to disappear from the fossil record in southwestern
Western Australia (Prideaux et al., 2010). Direct uranium series-
electron spin resonance (US-ESR) dating on megafauna tooth
enamel from several sites in South Australia (Grün et al., 2008),
from Cuddie Springs in New South Wales (Grün et al., 2010), and
from Titan’s Shelter in Tasmania (Cosgrove et al., 2010), also found
no extinct taxa younger than the Roberts et al. (2001) extinction
window, but a precise timing for the extinctions remains elusive.
Our direct 14C dating confirms that Protemnodon survived at Mt
Cripps until w41 ka cal BP, and adds a second extinct taxon, S.
occidentalis, to the megafauna of Tasmania with reliable ages
<50 ka cal BP. Many of the Mt. Cripps karst area cave deposits,
including CP213 and CP222, contain in situ articulated or associated
skeletons, indicating that little or no post-depositional reworking
has occurred, and when taken as a whole, record faunal change
across the megafaunal extinction boundary, with caves in the area
still acting as pitfall traps for the modern fauna (Camens, unpubl.
data). For cave CP213, where there is essentially no stratigraphy, our
directly-dated surface finds show that three extant species died in
the same ca. 45e41 ka cal BP period as two extinct Protemnodon
individuals.
4.2. Archaeological sites
The reliability of radiocarbon dating for many mainland
Australian archaeological sites has also been disputed (e.g. Allen
and Holdaway, 1995; Chappell et al., 1996; Bowler et al., 2003;
Gillespie et al., 2006; Field et al., 2008; Grün et al., 2010). Some of
the “chronometric hygiene” principles proposed by Spriggs (1989)
have been applied to the 14C record from Tasmanian rock shelter
archaeological sites by Holdaway and Porch (1996), eliminating
results with larger than usual error bars, non-finite ages, and ages
clearly inconsistent with others in the same site sequence; the
acceptable screened ages are listed in Table 3 and graphed in Fig. 4.
The oldest published Tasmanian archaeological sites have cali-
brated 14C ages on charcoal of 39.9  1.2 ka cal BP from Warreen
Cave (Allen, 1996) and 38.7  1.4 ka cal BP from Parmerpar Meet-
haner (Cosgrove, 1995), while charcoal from archaeologically sterile
layers has been dated to 39,970  950 BP at Parmerpar Meethaner
and at Parawee to >42,000 BP (Cosgrove et al., 2010). These authors
suggest that those dates provide evidence that humans were not in
 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47
Tasmania before 40 ka, in fact they show only that humans were not
at those sites before w44 ka cal BP. It is also possible that older
archaeological sites became submerged when the Bassian land-
bridge was flooded w14 ka (Lambeck and Chappell, 2001), as
previously suggested for mainland Australia (e.g. Chappell, 1993;
Oppenheimer, 2009).
Cosgrove et al. (2010) reported that “analysis of over 950,000
bones from the eight well dated and stratigraphically controlled
late Pleistocene archaeological sites from western, central and
northern Tasmania have not identified megafauna as part of human
prey.” However, this large number is for recovered bone fragments,
not whole bones, and since many of the bones were apparently
smashed to extract the nutritious marrow (Garvey, 2011), the
number of fragments from any one skeleton is also large. Very few
of these bone fragments are from the deepest layers of the oldest
sites, where the presence of extinct taxa might perhaps be antici-
pated. From Warreen Cave, for example, Cosgrove and Allen (2001)
estimated the minimum number of individual marsupials in the
oldest layers, ca. 35e40 ka cal BP, at three Macropus rufogriseus and
one V. ursinus. Allen (1996) notes that his excavation was “little
more than a test pit” with all the limitations implied by “telephone
booth” archaeology, and the directly-dated extant fauna (Cosgrove
et al., 2010) are significantly younger than the oldest charcoal.
Among the six archaeological sites with ages in excess of
30 ka cal BP included here, mostly excavated and dated more than
15 years ago, only one charcoal sample from Warreen Cave and one
from Parmerpar Meethaner have 2s age ranges extending older
than 40 ka cal BP. Similarly, only two Protemnodon bones from Mt
Cripps and two M. giganteus bones from Titan’s Shelter have 2s age
ranges extending younger than 41 ka cal BP (see Fig. 4). This is
a very small sample of both archaeological and fossil cave sites with
ages relevant to the contentious arguments about megafauna
extinction. Cosgrove et al. (2010) conclude that the archaeological
record implies that the Tasmanian megafauna were rare, or that
humans did not hunt them, or that the extinctions happened prior
to human arrival in Tasmania. We contend that the third possibility
has now been ruled out.
We dismiss the oft-repeated claims for late-surviving mega-
fauna at several archaeological sites in greater Australia. Cuddie
Springs in northern New South Wales, for example, has been widely
promoted as the only site on mainland Australian where extinct
fauna and archaeology occur in the same stratigraphic layers (e.g.
Field et al., 2008; Fillios et al., 2010), but the large scale excavations
and multiple dating methods applied do not support the conclusion
that bones and stones are the same age. Cuddie Springs has
previously been rejected because (a) single-grain OSL dating of
sand found multiple age distributions with maxima in two strati-
graphic units ca. 30 and 36 ka (Roberts et al., 2001), (b) statistical
Mt Cripps & Warreen Cave
Extant (n=6)
Titan’s Shelter
M. giganteus (large morph, n=4)
Mt Cripps Extinct (n=8)
50
Fig. 5. Summed probability distributions for calibrated ages measured on Tasmanian extant and extinct marsupials, and on archaeological charcoal, colours as for Fig. 4, with
w43 ka age for the Bassian landbridge emergence (Lambeck and Chappell, 2001). Fossil data from this work, Turney et al. (2008) and Cosgrove et al. (2010), archaeological data as in
Table 3, generated with OxCal 4.1 (Bronk Ramsay, 2009) using the IntCal09 atmospheric dataset (Reimer et al., 2009); all plots have the same vertical scale, baselines were shifted for
clarity.
45
analysis of published 14C ages on charcoal found that all samples in
the same two units had an error-weighted mean age of
35.9  2.2 ka cal BP with no ageedepth relationship (Gillespie and
Brook, 2006), and (c) detailed geochemistry and direct US-ESR
dating (Grün et al., 2010) found that the sand, charcoal and
extinct megafauna remains cannot be in primary context with their
ages represented by the OSL ages. We support the geochronology
rather than archaeological speculation, contra Field et al. (in press),
and reject Cuddie Springs as an extinct megafauna-human overlap
site. Nombe Rockshelter in Papua New Guinea (Gillieson and
Mountain, 1983), Cloggs Cave in Victoria (Flood, 1973), Seton
Rockshelter on Kangaroo Island, South Australia (Hope et al., 1977),
and the Tedford Locality at Lake Menindee in western New South
Wales (Cupper and Duncan, 2006) are also removed from conten-
tion, because they have no directly-dated extinct fauna and the
inclusion of older fossils in the dated archaeological horizons
cannot be precluded.
4.3. Potential bias in sample selection and site conditions
The small number of well-dated sites may suggest a tapho-
nomic or sampling bias in the Australian fossil and archaeological
records. Recent modelling studies support the idea that younger
ages tend to be more common in datelists because “the longer it is
since something happened, the greater is the probability that
evidence of that event will have disappeared” (Johnson and Brook,
2011). This bias in some measure represents the combined effects
of site erosion, taphonomy, diagenesis, site destruction and
opportunistic sampling, which may be detected (and potentially
corrected for) by comparing archaeological and geological dating
results (e.g. Surovell et al., 2009). All of the Tasmanian archaeo-
logical and fossil sites with acceptable ages listed in Table 3 and
graphed in Fig. 4 are caves or rockshelters; the fossil sites are
pitfall traps, the archaeological sites presumably not. From their
analysis of 822 Holocene 14C ages from 267 Australian rockshelter
sites, Johnson and Brook (2011) suggest that “.if the evidence
dated by archaeologists is more easily erased than the material
used to date geological substrates, the taphonomic effect on
archaeology will be underestimated.” For the much older sites
relevant to the Australian late Pleistocene extinctions, contra
Cosgrove et al. (2010), the absence of extinct taxa in known
Tasmanian archaeological deposits may only suggest that sites
from the earliest human occupation of Tasmania have yet to be
identified or adequately sampled. We also note that large uncer-
tainties are given for the ages and sill elevations used by Lambeck
and Chappell (2001) to derive the w43 ka Bassian landbridge
emergence time, which may change with new data (J. Chappell,
personal communication 2011).
Bassian Landbridge
Archaeological charcoal (n=16)
45 40 35 30
Calibrated Radiocarbon Age (ka cal BP)
46 R. Gillespie et al. / Quaternary Science Reviews 37 (2012) 38e47
4.4. Summary of the Tasmanian evidence
There are clear parallels between the few well-dated fossil and
archaeological records from the earliest period of human occupa-
tion on mainland Australia with the even smaller number of Tas-
manian records discussed here. The graph in Fig. 5 shows the 2s
summed probability distributions for calibrated ages measured on
Tasmanian extant fauna (n ¼ 6), large morph M. giganteus (n ¼ 4),
extinct megafauna (n ¼ 8) and archaeological charcoal (n ¼ 16),
generated with OxCal 4.1 (Bronk Ramsay, 2009); the vertical scale is
identical for each plot, with baselines adjusted for clarity. Although
they may not quite touch spatially, there is temporal overlap of
humans with the extinct P. anak and a large Pleistocene morph of
the extant M. giganteus.
5. Conclusions
The work reported here on fractions prepared from marsupial
bones and teeth from Mt Cripps, Tasmania, suggests that variations
of the Oxford ultrafiltration protocol yield acceptable radiocarbon
ages on well-preserved skeletal remains from Tasmanian sites near
the limits of the radiocarbon method. Our attempts to obtain reli-
able ages using the same methods on contaminated bones from
Mowbray Swamp were unsuccessful, and we conclude that an
atomic C:N ratio in the guideline 2.9e3.5 range is a necessary but
not sufficient quality control criterion.
No sites containing the coeval remains of both human occupa-
tion and directly-dated extinct megafauna have been documented,
and no human-extinct taxon interaction has been unequivocally
demonstrated, anywhere in greater Australia. However, the data
shows that some of the Mt Cripps now-extinct P. anak individuals
dated in this study and by Turney et al. (2008), as well as some of
the Titan’s Shelter extant M. giganteus (large morph) individuals
dated by Cosgrove et al. (2010), were alive w41 ka cal BPdafter
humans could first have walked across the emerging Bassian
landbridge from mainland Australia to Tasmaniadand therefore
the potential for human-extinct taxon interaction did exist.
Furthermore, we confirm that a second extinct megafaunal taxon, S.
occidentalis, was present w48 ka cal BP at Mt Cripps, and given that
there is no evidence of this species being in decline prior to human
arrival in Tasmania, it likely can also be counted among potential
interactees.
These geochronological results are closing the temporal gap
separating the archaeological record from the natural record of
megafauna in Tasmania, and we speculate that such investigations
will reveal the first convincing evidence of human interaction with
megafauna in Australia in the not too distant future.
Acknowledgements
We thank P. Darby and L. Gray for guiding us through the Mt.
Cripps karst, for assistance in the field we thank J. Metcalf, B.
Llamas, J. Wood, F. Salt, H. Geraldene and I. Houshold, and for
assistance in identifying specimens we thank G. Medlin, G. Pri-
deaux and W. Boles. For access to reference specimens we thank M.
Binnie and B. McHenry at the South Australian Museum, and L.
Gershwin at QVMAG. We appreciate the assistance provided by A.
Vains in sampling bones from the QVMAG collections, G. Jacobsen
for access to laboratory facilities at ANSTO, and T. Higham for the
Oxford data. For useful discussions we thank R. Cosgrove, E. Col-
houn, D. Bowman, B. Murphy, C. Johnson, R. Roberts and J. Chappell.
Funding for RG’s travel to Launceston was provided by the School of
Earth and Environmental Sciences, University of Wollongong, and
14
C dating at ANSTO was funded by AINSE grants 09/068 and 10/143
to RG (University of Wollongong). Funding for field work at Mt
Cripps and 14C dating at Oxford was provided by ARC Discovery
grant 20101413 to AC (Australian Centre for Ancient DNA).
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