About Journal of the North Atlantic
Isotopic Analysis of Faunal Material from South Uist, Western Isles,
Scotland
Jacqui Mulville, Rich Madgwick, Rhiannon Stevens, Tamsin O’Connell, Oliver Craig, Adrienne Powell,
Niall Sharples, and Mike Parker Pearson
Journal of the North Atlantic, Volume 2 (2009): 51–59
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Isotopic Analysis of Faunal Material from South Uist, Western Isles,
Scotland
Jacqui Mulville1,*, Rich Madgwick1, Rhiannon Stevens2, Tamsin O’Connell2, Oliver Craig3, Adrienne Powell1,
Niall Sharples1, and Mike Parker Pearson4
Abstract - This paper reports on the results from stable isotope analysis of faunal bone collagen from a number of Iron Age
and later sites on the island of South Uist, in the Western Isles, Scotland. This preliminary investigation into the isotopic
signatures of the fauna is part of a larger project to model the interaction between humans, animals, and the broader environment
in the Western Isles. The results demonstrate that the island fauna data fall within the range of expected results
for the UK, with the terrestrial herbivorous diets of cattle and sheep confi rmed. The isotopic composition for pigs suggests
that some of these animals had an omnivorous diet, whilst a single red deer value might be suggestive of the consumption
of marine foods, such as by grazing on seaweed. However, further analysis is needed in order to verify this anomalous
isotopic ratio.
1School of History and Archaeology, Cardiff University, Humanities Building, Colum Drive, Cardiff, UK CF10 3EU.
2McDonald Institute for Archaeological Research, Cambridge University, Downing Street, Cambridge, UK CB2 3ER.
3Department of Archaeology, The Kings Manor, York, UK Y01 7EP. 4Department of Archaeology, University of Sheffi eld,
Sheffi eld, UK S10 2TN. *Corresponding author - mulvilleja@cardiff.ac.uk.
Introduction
This paper presents results from a preliminary
study into faunal isotopic signatures from Prehistoric
and Norse sites on the island of South Uist in
the Western Isles of Scotland. The principal aims of
the investigation are to reconstruct changing foddering
regimes for the primary domesticates and
dietary strategies among red deer in this restricted
marginal island environment. This study represents
one line of evidence in a broader project to model
the interaction between humans, animals, and the
wider environment in the Western Isles (e.g., Smith
and Mulville 2004) based on data from the numerous
excavations undertaken by the Sheffi eld Environmental
Archaeological Research Campaign in the
Hebrides (SEARCH) and allied projects.
The ratio of the stable isotopes of carbon (δ13C)
and nitrogen (δ15N) in bone collagen provides an established
method for reconstructing the protein part
of the diet of past populations (Ambrose 1993, 2000;
Katzenberg 2000; Sealy 2001). The δ13C values give
an indication of the proportional contribution of terrestrial
and marine sources of dietary protein, whilst
δ15N values refl ect trophic level. In environments
that lack C4 plants, such as temperate Europe, high
δ13C collagen values demonstrate a contribution of
marine-derived protein to the diet (e.g., Barrett et
al. 2001), whilst high δ15N values are indicative of
the consumption of products from higher in the food
chain (e.g., meat, fi sh, or dairy protein) (Schoeninger
et al. 1983).
The majority of previous archaeological isotopic
studies have focused on reconstructing the
diet of human populations, but recent research
has recognised the importance of faunal isotopic
composition in providing a baseline from which
to interpret human collagen data (e.g., Hedges and
Reynard 2007, Jay and Richards 2007, Kosiba et
al. 2007, O’Connell and Kimball 2006, Privat et al.
2002, Richards et al. 2006). Climatic changes can
affect fractionation within the carbon and nitrogen
cycles and result in small-scale variation in isotope
signatures (Drucker et al. 2003, Murphy and Bowman
2006, Richards and Hedges 2003, Stevens and
Hedges 2004, Stevens et al. 2008, van Klinken et
al. 1994), though more local environmental effects
may also occur, such as elevated nitrogen isotope
values from sea spray (Britton et al. 2008, Virginia
and Delwiche 1982). Consequently, samples from a
range of food species are critical for the interpretation
of human isotopic signatures and for modelling
the interactions between humans, animals, and the
environment over time.
The technique also has the potential to inform
on human-mediated changes in animal diet, such as
changes in the exploited pasturage, manuring practices
(Bogaard et al. 2007, Jay and Richards 2006),
and fodder provision at a population and individual
level (e.g., the use of seaweed as fodder; Balasse et
al. 2005, 2006). The identifi cation of alternative fodder
resources are of particular relevance to insular
archaeology where high levels of neonatal mortality
in cattle have been attributed to the low availability
of hay fodder (Mulville et al. 2005). The exploitation
of a diverse range of terrestrial and abundant marine
resources, both directly and indirectly (as evidenced
by the zooarcheaological and palaeobotanical record;
Smith and Mulville 2004) results in complex
human-mediated ecosystems that are challenging to
model. The wide chronological range and quality of
the archaeological record in the Western Isles marks
these sites out as an excellent choice for examining
2009 Journal of the North Atlantic 2:51–59
52 Journal of the North Atlantic Volume 2
both cultural and environmental effects on stable
isotope values.
For this study, isotopic analysis was carried out
on a new sample of Late Iron Age specimens from
the multi-period assemblage at Bornais (Sharples
2005), and these results are compared with existing
faunal data from other sites on South Uist (Bronze
Age Cladh Hallan [Parker Pearson et al. 2005, 2007]
and the later Norse assemblage at Bornais). Research
on the human remains from these sites is also
in progress (e.g., Parker Pearson et al. 2005, 2007),
but is not discussed here.
Background
The sites
The settlement at Bornais was established in the
Late Iron Age with the construction of a wheelhouse,
which burnt down and was re-built as a rectangular
building (4th to 6th century AD). This was succeeded
by a Pictish phase of occupation with the later construction
of up to 20 buildings in the Norse period
(10th to 14th century AD). Bornais represents one of
the largest and most important Norse settlements in
the North Atlantic (Sharples 2005). Cladh Hallan is
an Early Bronze Age to Iron Age settlement lying
about 5 km south of Bornais and consisting of a
number of roundhouses in its later phases. The fi rst
of these roundhouses was built around 1300 BC, and
this long-lived site was occupied until around 500
BC (Parker Pearson et al. 2005, 2007).
Methods
Data are presented here for 40 Late Iron Age faunal
specimens from Bornais, analysed in 2007 at the
McDonald Institute, Cambridge, UK by the authors.
Also presented are additional results from Late Iron
Age (n = 14) and Norse Bornais (n = 21) obtained from
radiocarbon samples analysed at the Scottish Universities
Environmental Centre (SUERC) or the Oxford
Radiocarbon Accelerator Unit (ORAU) as part of the
site-dating program. The Cladh Hallan values were
obtained from a specifi c study of isotopic signatures at
the site undertaken at Newcastle University.
The 2007 samples were selected on a number of
criteria: to include a range of terrestrial species, to
avoid duplication of results from any one individual,
and to avoid age-related dietary changes. Samples
from three domestic (cattle [Bos Taurus], sheep
[Ovis aries], and pig [Sus scrofa]) and one wild
species (red deer [Cervus elaphus]) were selected
and, when possible, separate animals were identifi
ed (e.g., by repeatedly sampling the same part and
side of a particular element). Aging information
was utilized to ensure that juveniles below weaning
age were omitted. Collagen from the 2007 material
was extracted following a modifi ed Longin method
(Longin 1971). Faunal samples were selected for
the radiocarbon-dating programs without particular
regard to the avoidance of repeat samples of single
individuals or juveniles.
For all of the samples, material was analysed
using an automated elemental analyzer coupled in
continuous-fl ow mode to an isotope-ratio-monitoring
mass-spectrometer (in Cambridge, a Costech
elemental analyser coupled to a Finnigan MAT253
mass spectrometer, and in Oxford/Newcastle, a
Carlo Erba elemental analyzer coupled to a PDZ
Europa Geo 20/20 mass spectrometer). Stable isotope
concentrations are measured as the ratio of the
heavier isotope to the lighter isotope relative to an
internationally defi ned scale—VPDB for carbon,
and AIR for nitrogen (Hoefs 1997). Isotopic results
are reported as δ values (δ13C and δ15N) in parts per
1000 or “per mil” (‰) values, where
δ15NAIR = ([15/14Nsample / 15/14NAIR] - 1) × 1000.
Based on replicate analyses of international and laboratory
standards, measurement errors are less than
±0.2‰ for δ13C and δ15N. The analytical errors on
samples analysed at ORAU are larger, potentially as
large as ±0.4‰ (Peter Ditchfi eld, Research Laboratory
for Archaeology and the History of Art, University
of Oxford, Oxford, UK, pers. comm.). No inter-laboratory
repeat testing was undertaken, but all samples
were measured relative to International Atomic Energy
Agency standards, with the McDonald Institute
and ORAU also measuring the samples relative to the
same in-house standard. Thus, the results from different
laboratories are thought to be comparable.
Results
The Western Isles
Faunal isotope data for the sites are presented
in Table 1. The Bornais Late Iron Age samples all
yielded collagen and had atomic C/N ratios of 3.2 to
3.3, well within the range deemed to be indicative of
acceptable collagen preservation (Ambrose 1990).
The mean δ13C results from the combined Western
Isles sites for cattle (-21.4‰), sheep (-21.1‰),
and red deer (-21.5‰) samples are typical for herbivores
living in a temperate C3 ecosystem such
as Britain (Richards and van Klinken 1997, van
Klinken et al. 2000), and do not indicate the consumption
of significant amounts of marine dietary
sources. The mean δ15N values for the herbivorous
species from the South Uist sites—cattle (4.8‰),
sheep (4.6‰), and deer (4.4‰) values—are typical
of those found elsewhere for UK herbivores
during the Holocene. The pig values, based almost
exclusively on Late Iron Age animals with only a
single Norse individual, differ in δ15N compared to
those of the cattle and sheep, with average values
of -20.4‰ (δ13C) and 7.1‰ (δ15N). There is a small
2009 J. Mulville, R. Madgwick, R. Stevens, T. O’Connell, O. Craig, A. Powell, N. Sharples, and M.P. Pearson 53
trophic-level increase in faunal δ13C (0 to 2‰)
and δ15N (3 to 5‰) values (e.g., Bocherens and
Drucker 2003, Rau et al. 1983, Schoeninger 1985,
Schoeninger and DeNiro 1984), and carnivores
and omnivores have higher δ13C and δ15N values
than herbivores (Roth and Hobson 2000). These
increased values of δ13C and δ15N in pigs may result
from variation in the isotopic signal of the plant
foods consumed (e.g., seeds vs. forage), or some
animal protein input, such as scraps of meat, fish,
dairy protein, or excrement. The range of variation
within species (cattle 2.0‰, sheep 1.8‰, pig
1.8‰, red deer 2.5‰) is similar to that seen at other
archaeological sites with large suites of faunal analyses,
e.g., Neolithic Hazleton (Hedges et al. 2008).
Chronological changes
The mean values for the individual species from
Bronze Age Cladh Hallan and the various phases at
Bornais can be compared across time, as presented
in Table 1 and Figure 1, with individuals of different
species plotted by site in Figure 2.
Table 1. Mean δ13C and δ15N values for different taxa from sites
on South Uist, Scotland.
Std Std
dev dev
δ13C δ15N δ13C δ15N n
Cladh Hallan Bronze Age
Cattle -21.7 5 0.5 0.6 12
Sheep -20.1 5.9 0.4 0.6 2
Red Deer -21.2 4.9 0.0 0.5 2
Bornais Late Iron Age
Cattle -21.1 4.3 0.4 0.5 15
Sheep -21.3 4.4 0.3 0.6 12
Pig -20.6 6.9 0.4 0.8 9
Red Deer -21.7 4.4 0.4 0.7 18
Bornais Pictish/Norse
Cattle -21.4 5.1 0.5 1.6 16
Sheep -21.2 4.6 0.6 1.2 3
Pig -19.3 8.7 - - 1
Red Deer -19 4.1 - - 1
Western Isles Combined
Cattle -21.4 4.8 0.5 1.2 43
Sheep -21.1 4.6 0.5 1.3 17
Pig -20.4 7.1 0.6 0.9 10
Red Deer -21.5 4.4 0.7 0.7 21
Figure 1. δ13C and δ15N values from cattle, sheep, pig, and red deer samples from South Uist, Scotland by site.
54 Journal of the North Atlantic Volume 2
Figure 2. δ13C and δ15N values from cattle, sheep, pig, and red deer samples from South Uist, Scotland by species.
2009 J. Mulville, R. Madgwick, R. Stevens, T. O’Connell, O. Craig, A. Powell, N. Sharples, and M.P. Pearson 55
Cattle have similar δ13C values (-20.5‰ to
-22.5‰) through time, but vary slightly in δ15N,
with the Iron Age values lower compared to the
other two sites. There are two outliers with higher
than expected δ15N values (8.6‰ and 9.6‰) from
the Norse period. These values were obtained from
radiocarbon samples, and in the absence of detailed
aging, may represent unweaned calves (Balasse and
Tresset 2002, Hedges and Reynard 2007, Jenkins
and Partridge 2001). A third outlying Norse cattle
specimen has a low δ13C level (-22.7‰).
The Bronze Age sheep at Cladh Hallan have
slightly higher δ15N and δ13C values than the later
sites. In such a small sample, the variation between
these two sites is not substantial enough to be indicative
of radically different feeding strategies or wider
climatic changes. The few Norse sheep lie close to
the range for Iron Age sheep.
Figure 2 demonstrates that the Bornais Iron Age
pigs values, with averages of -20.6‰ (δ13C) and
6.9‰ (δ15N), differ markedly from cattle and sheep
values, with the single Norse animal shifted even
further along the δ13C axis. This shift in δ13C and
δ15N relative to the herbivorous cattle and sheep
in the late Iron Age pig sample is indicative of a
diet with an input from animal protein and possibly
some marine foods. The Bronze Age and Norse
datasets need considerable expansion before firm
conclusions can be drawn on chronological shifts in
pig diet.
With the exception of a single Pictish animal,
all red deer specimens fall within the range of the
domestic herbivore values. The Pictish red deer lies
outside this range; its δ13C value of -19‰ is higher
than all of the pig specimens. In comparison, a study
of 129 modern island British red deer indicated collagen
δ13C values ranging from -24.8‰ to -21.2‰
(Stevens et al. 2006), which is equivalent to -23.9‰
to -20.3‰ in archaeological populations accounting
for fossil-fuel effects (Friedli et al. 1986). The
elevated δ13C of this single deer could be indicative
of a marine element in feeding (Balasse et al. 2005,
2006). Modern red deer are known to graze upon
seaweed (Fig. 3; Conradt 1999), which may explain
the carbon shift in this archaeological specimen.
The wider archaeological context
Data for the main mammal groups from a range
of Iron Age sites across Scotland (Lismore, Inner
Hebrides [O’Connell and Kimball 2006]; and Port
Seaton, Broxmouth, and Dryburn Bridge, East Lothian
[Jay and Richards 2007]), the earlier Iron Age
sites of England (Wetwang Slack, Yorkshire [Jay
and Richards 2006]; Micheldever Wood and Winnall
Figure 3. Modern red deer stag grazing on seaweed on Rum (photograph © Deer Commission for Scotland).
56 Journal of the North Atlantic Volume 2
Down, Hampshire; Harlyn Bay and Trevelgue Head,
Cornwall [Richards and Jay 2007]; and Yarnton, Oxfordshire
[Lightfoot et al., in press]) and the Norse
site at Newark Bay, Orkney in the Northern Isles
(Richards et al. 2006) are presented in Table 2.
The Western Isle cattle and sheep δ13C and δ15N
values fall within the range found at the majority of
comparative sites. The δ13C values for herbivores
(cattle, sheep, and red deer) range from -22.7 to
-20.6‰, with the Lismore animals forming the
bottom end of the range (-21.9 to -22.7‰) and
occasional higher values scattered across the sites.
The δ15N values are more variable, with high sheep
δ15N values noted from Newark Bay, Broxmouth,
and Yarnton, and with cattle values also elevated
at the latter two. In a recent review of a number of
these sites, this variation in nitrogen values has been
attributed to variation in “baseline” environmental
values (from the plants) at different locations (Jay
and Richards 2007). At individual sites, this variation
has been attributed to different management
strategies, with animals grazing particular plants
(e.g., salt-marsh vegetation; Britton et al. 2008) and/
or a greater exposure to a sea-spray/salinity effect
(Heaton 1987, Richards et al. 2006). The data reviewed
in this paper does not support the existence
of similar mechanisms for δ15N enrichment at all
coastal environments (contra Britton et al. 2008).
Cattle and sheep values in the Western Isles and
Cornwall, and cattle values in East Lothian are relatively
low. Thus, high δ15N values in the Bronze Age
Seven Estuary (Britton et al. 2008) may be directly
related to the specifi c salt-marsh vegetation in particular
rather than to coastal vegetation in general.
The pig δ13C values at the comparative sites are
similar to those of the herbivores, but the range of
δ15N values are higher (5.7 to 9.2‰), suggestive of an
omnivorous diet at some of the sites, as has been described
for the Hebrides. This is not true of all sites,
and at Lismore, in East Lothian, the similar δ13C and
δ15N isotope values of the pig compared to sheep and/
or cattle are indicative of an herbivorous diet. Red
deer values are reported from four sites, all of which
show lower average δ13C values compared to sheep
and cattle (-21.9 to -22.1‰) and a narrow range of
δ15N values (4 to 5.4‰) that lie within the range reported
by Stevens et al. (2006). There is no evidence
for higher carbon isotope values comparable to those
reported from the Western Isles Norse deer and pig.
This result highlights the unusual nature of these individual
animal diets and that further research on the
possible input of marine foods is required.
Discussion
These initial results have highlighted a number
of points. Within the Western Isles, the representative
nature of the dataset needs to be confi rmed by
enlarging the sample sizes to cover the full range of
species across the chronological spread, to incorporate
aging information into the datasets, and to
avoid inter-laboratory biases. Data obtained from
radiocarbon determinations have provided useful
supplementary information, but slight measurement
differences between laboratories could be signifi -
cant in such small samples, especially when dealing
Table 2. Mean δ13C and δ15N values for different faunal taxa at a
selection of British Prehistoric sites.
Std Std
dev dev
Site δ13C δ15N δ13C δ15N n
Inner Hebrides
Lismore Iron Age
Cattle -22.4 4.6 0.5 0.6 5
Sheep/goat -22.7 6.0 0.4 0.9 10
Pig -21.9 6.0 0.7 0.8 4
Red deer -22.1 4.0 0.5 0.1 2
Northern Isles
Newark Bay Norse
Cattle -21.8 5.5 0.2 0.8 9
Sheep/goat -21.8 7.0 0.8 0.9 5
Pig -20.6 8.4 1.3 1.6 6
East Lothian
Broxmouth Iron Age
Cattle -21.9 6.1 0.2 0.7 5
Sheep/goat -21.6 6.4 0.2 0.8 7
Pig -21.8 7.7 0.4 0.4 6
Red deer -22.1 4.4 0.4 1.4 4
Dryburn Bridge Iron Age
Cattle -21.8 5.6 0.1 1.1 4
Sheep/goat -21.8 6.3 - - 1
Pig -21.3 6.8 - - 1
Port Seaton
Cattle -21.7 5.5 0.3 1.2 4
Sheep/goat -21.8 7.4 0.3 0.4 3
Pig -21.6 9.2 - - 1
Yorkshire
Wetwang Slack Iron Age
Cattle -21.5 4.6 0.5 1.1 10
Sheep/goat -21.6 4.9 0.6 1.0 17
Pig -21.6 6.4 0.6 1.5 11
Oxfordshire
Yarnton Iron Age
Cattle -21.6 6.9 0.3 1.4 12
Sheep/Goat -21.5 6.6 0.2 1.3 12
Pig -21.7 8.1 0.4 0.8 12
Hampshire
Micheldever Wood Iron Age
Pig -21.3 6.1 0.5 0.2 5
Winall Down
Cattle -21.8 4.5 0.4 1.1 11
Sheep/goat -21.7 4.2 0.6 0.8 9
Pig -21.4 5.7 0.6 1.0 4
Red deer -21.9 5.2 - - 1
Cornwall
Halyn Bay Iron Age
Cattle -21.9 5.8 0.1 0.0 2
Pig -22.4 5.7 - - 1
Trevelgue Head
Cattle -22 4.8 0.3 2.0 7
Sheep/goat -21.8 5.8 0.3 0.5 8
Pig -21.5 7.1 0.4 1.4 8
Red deer -22 5.4 0.2 0.5 4
2009 J. Mulville, R. Madgwick, R. Stevens, T. O’Connell, O. Craig, A. Powell, N. Sharples, and M.P. Pearson 57
with small isotopic shifts. The absence of pig and
red deer data within the radiocarbon samples from
Cladh Hallan and Norse Bornais, an indirect effect
of the lower abundance of these species, hampered
chronological comparisons.
The Western Isles results for domestic animals can
be placed in context by comparison with wild species,
as they are considered to provide a more accurate representation
of isotopic signals in the wider environment.
These species are free to make choices regarding
their food input, and for the purpose of this study,
red deer are assumed to refl ect the stable isotope
signal of freely grazing animals. The similarity in the
level and low range of isotopic composition for the
Western Isle deer and sheep suggests these animals
have similar, herbivorous feeding patterns. However,
there is a potential problem in using large wild herbivores
to provide “natural” signals as their behaviour
can be affected by humans—directly through hunting
and indirectly through habitat destruction and
disturbance which can promote changes in behaviour.
This infl uence may be a particular problem in the
Hebrides, where red deer play a large part in the food
economy (Mulville, in press). In addition, the presence
of anomalous values from a probable seaweedeating
deer highlights the variety that can be found
with wild faunal diets. Future work on stable isotopes
in the Western Isles will target additional wild species
to establish background isotopic signals.
Interspecies comparisons within this Western
Isles dataset demonstrate that the dietary signal for
pigs is different to that of the herbivorous animals,
with a distinctive animal protein input identifi ed in
the Late Iron Age Western Isles sample. The omnivorous
nature of Late Iron Age pig diets in the
Western Isles is consistent with the zooarchaeological
models for their husbandry, with the low incidence
of pig farming linked to the lack of available
fodder and the potential for destruction of the plant
cover by allowing these rooting animals to forage
freely, exposing the light, unconsolidated soils
to the strong winds characteristic of the Hebrides
(Serjeantson 1990). The occasional pig is thought to
have been reared by providing animal and plant food
sources which would include scraps of meat, fi sh,
and dairy waste along with excrement from humans
and animals (Mulville 2005, Serjeantson 1990). The
stable isotope data seem to confi rm this model of
small-scale intensive pig husbandry.
Overall, the results from the Western Isles lie
within the range of other reported Iron Age and Norse
sites (e.g., Jay and Richards 2007:187), despite their
special status in terms of environment, marginality,
and the extended duration of the Scottish Iron
Age. There is a demonstrable degree of variation
between the sites, particularly in nitrogen levels,
which echoes comments made by Jay and Richards
(2007), and suggests regional, chronological, and
environmental variation. Further work is needed to
elucidate the causes, particularly in relation to the
effects of differing vegetation and exposure to sea
spray. This study further reinforces the necessity for
a good faunal baseline against which to measure all
interpretations of human diet from carbon and nitrogen
isotopic compositions for each individual locale
(Jay and Richards 2007).
Conclusion
There are apparent isotopic differences between
the Western Isle datasets, between the Scottish island
sites, and also between the island and mainland sites,
but these differences seem unrelated to larger environmental
factors. However, although these results
provide potential indications of both inter- and intrataxon
variation in feeding patterns, sample sizes are
at present too small for confi dent interpretations to
be made, as these limited datasets may contain individuals
that are outliers to the general population. In
addition, temporal climatic changes that impact on
isotopic composition could be responsible for the
inter-site variation on the islands, and it remains diffi
cult to observe temporal diversifi cation in animal
management strategies. These results have been
successful in picking up inter-species differences,
which can be integrated with zooarchaeological
analyses. There is a diversifi cation of pig dietary
foods not found elsewhere, although evidence for
the use of marine foods as an additional source of
fodder for domestic cattle has not been proven.
The range of faunal values demonstrated within
and between the data sets continues to add to recent
debates on the validity of isotopic models and the
need for good faunal data upon which to base human
dietary reconstructions and to better understand
wider climatic shifts (Britton et al. 2008; Hedges and
Reynard 2007; Jay and Richards 2007; Stevens et
al. 2004, 2006). This study provides a starting point
for a comprehensive program of isotopic analysis on
larger samples from sites on the Western Isles and
represents only an initial step in the large-scale isotopic
modelling of a restricted island eco-system.
Acknowledgements
Thanks to Historic Scotland for funding the excavations
at Cladh Hallan and Bornais, the Red Deer
Commission for the use of the photograph in Figure 3, and
two anonymous reviewers for their insightful comments
on an earlier draft of this paper. Thanks to Mike Hall and
James Rolfe, Godwin Lab, Department of Earth Sciences,
Cambridge University for help with isotopic analyses and
the staff of SUERC and ORAU.
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