251ISSN 2227-4952 (Print), ISSN 2708-6143 (Online). Археологія і давня історія України, 2020, вип. 4 (37)
УДК 902.65(477) DOI: 10.37445/adiu.2020.04.20
M. C. Lillie, C. E. Budd 
DIET ISOTOPE ANALYSIS AND RELATED STUDIES  
IN PREHISTORIC UKRAINE: FACT, FICTION AND FANTASY 
This paper outlines the results of twenty-eight years 
of collaborations between the authors and colleagues 
in Kiev, initiated when the first author began PhD re-
search at Sheffield University under the supervision of 
the late Professor Marek Zvelebil in 1992. From the out-
set of this doctoral research Professor Dmitri Telegin, to 
whom this paper is dedicated, and Dr. Inna Potekhina, 
were fundamental not only to the success of the original 
research programme, but in terms of the considerable 
generosity, insight and friendship that was extended to 
the lead author as he navigated his way through the 
earlier Holocene parts of Ukrainian prehistory. The 
current study is as much a result of the work of the cur-
rent authors as it is of collaboration and collegiality of 
these colleagues.
The topics considered throughout this paper focus 
around the key observations and themes that have been 
developed since the research began. It also aims to high-
light those areas where inconsistencies occur, and where 
clarification is deemed warranted due to the activities 
of researchers who have failed to fully appreciate the 
nuances of Ukrainian prehistory and multi-discipli-
nary research agendas. It is apparent that, in light of a 
recent «gold rush» to claim ownership of the materials 
available in Ukraine, at prehistoric sites of all periods, 
there is clearly a need for a considered and careful ap-
proach to the data generated from dietary isotope and 
related studies. Furthermore, our research since the 
early 1990s has shown that misidentification of frag-
mentary or isolated bone in both primary and second-
ary contexts can lead to erroneous interpretations and 
occasional «flights of fancy». This paper will outline a 
number of the issues identified, and also explore issues 
around data use and representation in an attempt to 
offer some balance to discussions of prehistoric diet and 
chronology in Ukraine.
Keywords: Ukraine, prehistory, dietary isotope 
analysis, δ13C, δ15N, AMS dating, freshwater reservoir 
effects.
Introduction. Studies of diet using stable 
isotope analysis alongside dental and skeletal 
pathology can provide detailed insights into the 
nature of the subsistence strategies employed by 
prehistoric groups, and their past lifeways. Ini­
tial investigations in the 1990s, into the dietary 
pathways of the Dnieper-Donets culture, through 
analysis of skeletal remains from the Mariupol-
type cemeteries, showed that these groups placed 
an emphasis on the consumption of terrestrial 
animals and freshwater resources, alongside veg­
etal foodstuffs (the latter being less well reflected 
in isotopic studies that use bone collagen — see 
discussion below). Alongside palaeopathological 
and osteological studies of the human populations 
from the Dnieper cemeteries the research agenda 
has integrated AMS radiocarbon dating in an ef­
fort to provide an absolute chronological frame­
work to the cemeteries studied. As discussed be­
low, studies that fail to undertake such analyses 
run the risk of misinterpretation of the results 
from parallel studies of diet and pathology, and 
can, as will be highlighted, produce fundamental­
ly flawed interpretations and misrepresentation 
of the evidence for past human lifeways.
Our investigations to date have confirmed 
that across the prehistoric periods, from at least 
10,200 cal BC through to ca. 3500 cal BC (and 
probably beyond) the consumption of freshwater 
resources, including species of fish such as carp 
and pearl roach, formed an integral, and fun­
damental, part of subsistence strategies in the 
Dnieper region (Lillie et al. 2011; Potekhina et al. 
2014). It has also been possible to demonstrate 
the existence of a freshwater reservoir effect that 
is influencing the absolute (radiocarbon) dating 
of the individuals that are interred in the cem­
eteries of the Dnieper region, where freshwater © M. C. LILLIE, C. E. BUDD, 2020
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resources are shown to be integral to subsistence 
strategies. As such, the available absolute dat­
ing of these cemeteries, when undertaken on the 
human remains themselves, or on freshwater re­
sources, must be considered with caution when 
discussing the socio-cultural evolution of the pop­
ulations being studied (Lillie et al. 2009).
Importantly, the continued reliance on the ex­
ploitation of wild resources across the Mesolithic 
and Neolithic period’s means that the definition 
of the Neolithic does not necessarily follow the 
convention of a shift towards the exploitation of 
domesticates in many areas of Eastern Europe, 
as the transition lacks the major socio-economic 
transformations that generally demarcate the 
transition from a food extraction to a food produc­
tion economy (Lillie et al. 2020). Indeed, as previ­
ously noted by Dolukhanov and Khotinsky (1984), 
whilst the appearance of a production economy 
would normally indicate Neolithisation, other 
criteria, such as the first appearance of ceramics, 
improved working tools, new forms of hunting 
and an increased reliance on fishing are also used 
to define the transition to a «Neolithic» way of life 
in Ukraine, and many parts of Eastern Europe 
(also Motuzaitë Matuzevičiűtë, Telizhenko 2016). 
In addition, Telegin and Titova (1993) attributed 
all those cultures which are characterised by the 
presence of ceramics, but lacking metal, to the 
Neolithic period. Furthermore, it should also be 
noted that, given the areal extent of the region of 
study, it is perhaps unsurprising that the nature 
and timing of this transition displays significant 
asynchronicity when studied in both a west—east 
and a south—north direction across Ukraine (Lil­
lie et al. 2020, p. 188).
In general, research into the prehistoric popu­
lations of Ukraine has been developed within a 
rigorous methodological and scientific framework, 
but on occasion a lack of detailed knowledge and 
understanding of context has resulted in the 
publication of ill-conceived or poorly thought out 
research that has weakened the validity of new 
studies in this region (see Budd et al. 2020 for 
an overview of these studies). The current paper 
seeks to highlight areas where a more rigorous 
scientific and theoretical basis may be warranted, 
to consider aspects of recent studies undertaken 
by researchers who are relatively new to Ukrain­
ian prehistoric studies, and to outline areas that 
would benefit from a more considered approach, 
or from clarification of the results obtained; both 
in terms of context and previous studies (see cri­
tiques in Lillie 2020; Lillie et al. 2020).
1. background. The Dnieper Rapids region of 
Ukraine contains a wealth of cemeteries and bur­
ials dating from the earliest Holocene through to 
the later prehistoric periods and beyond (Telegin 
1986; Telegin, Potekhina 1987; Lillie 1996; 1998a; 
1998b). These cemeteries, the Dnieper-Donets 
Mariupol-type cemeteries (DD M-t), formed the 
basis of the first authors PhD research across the 
1990s, and more recently (over the past decade) 
has formed an integral element of the second au­
thors’ postgraduate and postdoctoral research at 
Oxford and Umeå Universities. Across this re­
search period a number of reassessments of the 
chronological development of the cemeteries and 
the socio-cultural developments in evidence have 
been undertaken in collaboration (e. g. Potekhina, 
Telegin 1995; Lillie 1998a; 1998b; 1998c; Telegin 
et al. 2000; 2002; 2003a; 2003b; Lillie et al. 2009; 
Budd, Lillie 2020), and an overview of this work 
is presented in our most recent collaboration (Lil­
lie et al. 2020).
Interpretations into the significance of the con­
centration of cemeteries at the Dnieper Rapids, 
and along the Dnieper itself, have reinforced ob­
servations relating to the importance of a reliable 
resource base to earlier Holocene fisher-hunter-
foragers in a region where the areas away from 
the river valleys are poorly watered and lack the 
rich vegetation niches found in the river valleys 
themselves. Dolukhanov and Khotinskiy (1984, 
p. 319) have previously argued that Mesolithic 
groups exploited the fauna of the steppe, for­
est and tundra zones, with fishing and gather­
ing becoming increasingly important factors in 
their adaptation to the more differentiated early 
Holocene landscapes (also Gimbutas 1956). In ad­
dition, Lillie (2003a, p. 4) has also suggested that 
the focus of large-scale cemeteries at the Dnieper 
Rapids argues for, at least, a seasonal aggrega­
tion of population at this location in order to ex­
ploit the stable resources, such as freshwater and 
anadromous fish, along with the plants and fauna 
of the riparian zone. Such aggregations would ob­
viously offer the opportunity for a wide range of 
socio-political as well as economical interactions 
between the various groups in and around the 
Dnieper and its tributaries, and this aspect of in­
ter-group interactions should not be overlooked.
At the cemeteries in this region the burial in­
ventories themselves, and finds from archaeologi­
cal sites, provide evidence that reinforces the im­
portance of fishing through the presence of fishing 
related artefacts such as harpoons, net sinkers, 
fish-hooks, and fish-tooth necklaces, which are, 
by their very nature, more likely to be reported 
(e. g., Telegin 1986; Telegin, Potekhina 1987; Lil­
lie, Richards 2000; O’Connell et al. 2000). The 
archaeological evidence from Neolithic and Eneo­
lithic sites in the Dnieper region indicates that 
the range of faunal and freshwater resources 
available during these periods, as shown by the 
finds from the Neolithic settlements of Sobachky, 
Sredny Stog I and Buzky (Telegin, Potekhina 
1987), and the Eneolithic site of Dereivka II (Tel­
egin 1986), included a broad range of both wild 
and also some domesticated species. For the Neo­
lithic period these included aurochs, red and roe 
deer, wild pig and beaver, alongside domesticates 
such as cattle, sheep / goat, pig, horse and dog 
(Dolukhanov 1979; Mallory 1987). In most cases 
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Lillie, M. C., Budd, C. E. Diet Isotope Analysis and Related Studies in Prehistoric Ukraine: Fact, Fiction and Fantasy
however, the evidence does not support the signif­
icant integration of domesticated species within a 
production economy, and instead the data prob­
ably suggest increasing trade and exchange be­
tween the fisher-hunter-foragers in the Dnieper 
region and adjacent groups of pastoralists and / 
or farmers.
Freshwater fish species such as European cat­
fish, perch, roach, rudd, asp and carp, along with 
European pond terrapin and otter, as well as 
waterfowl such as mallard, pintail duck, goose, 
teal and coot are all in evidence at the Eneolithic 
site of Dereivka (Telegin 1986, p. 84). In addition, 
freshwater mussel and river snails are also at­
tested. It should be anticipated that a similarly 
broad range of resources would have been avail­
able in earlier periods.
Whilst the archaeological data (e. g. Dolukhanov 
1979; Telegin 1986; 1987; Telegin, Potekhina 
1987; Kozłowski 1989) attests to the exploitation 
of a range of faunal and freshwater species, only 
limited research into palaeopathological analysis 
or dietary studies were undertaken prior to the 
work of Jacobs (1993; 1994) and Lillie (1996; 1997; 
1998a). Research by Lillie (1996; 1998a), initially 
aimed at characterising the transition to the Neo­
lithic from the perspective of palaeopathological 
analysis, soon identified the need for enhanced 
chronological studies (as discussed above), and 
integrated stable isotope studies through an ini­
tial collaboration with Mike Richards at Oxford 
University (Lillie, Richards 2000). The studies 
that have been undertaken since this early stage 
of the research agenda are summarised below.
2. Dietary Isotope Studies. Since the initial 
integration of dietary isotope studies into the re­
search agenda a number of papers have sought 
to expand the dataset through whole cemetery 
analysis and / or through integrated analysis of 
materials, whether from human, faunal or fish 
remains, as they became available (e. g. Lillie et 
al. 2003; 2011; 2016; Lillie, Jacobs 2006; Lillie, 
Budd 2011; Potekhina et al. 2014; Budd, Lillie 
2020; Budd et al. 2020).
Diet isotope studies have their limitations of 
course, and as noted in Lillie and Budd (2011, 
p. 44), there are differences in the δ13C values 
obtained when using the different tissues, for 
example bone collagen or apatite-carbonate, and 
consequently it is also important to note that 
the stable isotope analysis of bone collagen pre­
dominantly reflects the protein component of 
the diet (Krueger, Sullivan 1984; Ambrose, Norr 
1993; Tieszen, Fagre 1993; Schulting, Richards 
2001). As many plant foodstuffs are difficult to 
«see» isotopically because the protein levels of 
unprocessed plants are usually quite low when 
compared to meat (from fauna and fish), there is 
an inherent «bias» towards meat proteins when 
using bone collagen in isotope studies, as opposed 
to reflecting the diet as a whole (Müldner, Rich­
ards 2005 and references therein). However, de­
spite the fact that diet isotope studies using bone 
collagen are not ‘whole diet’ indicators, stable 
isotope analysis does provide a direct measure 
of the nature of past human diet, as the carbon 
isotope value (δ13C), indicates the amount of ma­
rine protein in the diet, as compared to terres­
trial protein, and bone collagen analysis also dis­
tinguishes between different dietary components, 
such as C3 and C4 photosynthetic plants and the 
animals that consumed them (Schwarcz, Schoe­
ninger 1991; Richards 2002). Nitrogen stable iso­
tope ratios (δ15N) are used to establish the trophic 
level of an organism in the food web, with δ15N en­
richment of c. 3—6 ‰ (e. g. Δ15Ndiet-body) observed 
as one progresses up the food chain (Schoeninger, 
DeNiro 1984; Minagawa, Wada 1984; Hedges, 
Reynard 2007).
Throughout the research programme the sam­
ples for diet isotope studies have been processed at 
the Research Laboratory for Archaeology and the 
History of Art, University of Oxford. In the initial 
studies (e. g. Lillie, Richards 2000; Lillie, Jacobs 
2006) the collagen was extracted from the human 
bone samples following the protocol outlined in 
Richards (1998). In the more recent work (Budd 
2015; Lillie et al. 2011, p. 61) the bone collagen 
was obtained from the skeletal material using a 
modified version of the «Longin» (1971) extrac­
tion method. Samples are subjected to a number 
of acid and H2O (MilliQ) washes and freeze dried. 
The resulting collagen is then analysed using an 
automated carbon and nitrogen analyzer and a 
continuous-flow isotope ratio-monitoring mass-
spectrometer (Carlo Erba carbon and nitrogen el­
emental analyser coupled to a Europa Geo 20/20 
mass-spectrometer). Typical replicate measure­
ment errors are of the order of ±0.2 ‰ for δ13C 
and δ15N. Only samples which produced collagen 
yields with a C:N ratio between 2.9 and 3.6 (De­
Niro 1985) were used in the analysis of palaeodi­
et. The following sections summarise the current 
situation with regards to the analyses that have 
been undertaken to date, the most recent updates 
for which appear in Budd and Lillie (2020), and 
Budd et al. (2020).
2.1. epipalaeolithic and mesolithic Sub-
sistence. The cemeteries and sites analysed in 
the investigation of earlier Holocene subsistence 
practices include human remains from Vasilyev­
ka III and II, Marievka and Osipovka, and the 
Crimean site of Fat’ma Koba. It also includes 
fauna remains from the Epipalaeolithic sites of 
Vasilyevka III, Voloshkoe and Vyazivok, and the 
Mesolithic sites of Fat’ma Koba and Rogalik 2 
(fig. 1). Research by Lillie and co-workers e. g. Lil­
lie (2003a), Lillie et al. (2003), Lillie and Jacobs 
(2006), Lillie et al. (2011) has previously sug­
gested that during the Epipalaeolithic and Me­
solithic periods the slightly higher δ15N values at 
the later Mesolithic site of Vasilyevka II (13.40 ± 
0.65 ‰ (at 1σ)), when compared to 12.75 ± 0.61 ‰ 
from Vasilyevka III, is suggestive of either a 
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higher input from freshwater resources in the 
later Mesolithic period, or alternatively, the ex­
ploitation of differing fish species from a range 
of trophic levels in the riverine ecosystem (Lillie 
et al. 2011, p. 63). In addition, the δ13C values at 
Vasilyevka II (–20.94 ± 0.51 ‰) are depleted by 
2 ‰ when compared to Vasilyevka III (–22.22 ± 
0.24 ‰; ibid. 2011, p. 61—63). Importantly, one 
individual of Mesolithic date from the predomi­
nantly Neolithic cemetery of Dereivka I, individ­
ual 84, also exhibits δ13C and δ15N values that are 
fully commensurate with the mainstream values 
evident through the Epipalaeolithic and Meso­
lithic periods at (–23.6 ‰ and 11.4 ‰, although it 
is noted that the δ15N value is towards the lower 
end of the ranges reported.
In this context, the data has shown that at the 
site of Marievka burials 4 and 14 have δ15N ratios 
that would be commensurate with a diet wherein 
the majority of dietary proteins are obtained from 
terrestrial as opposed to freshwater resources. As 
noted by Budd and Lillie (2020, p. 291) on the 
basis of the dating at Marievka, these two indi­
viduals represent the chronologically earliest evi­
dence for the exploitation of terrestrial as opposed 
to aquatic dominated diets. Unfortunately due to 
the limited number of interments at Marievka, 
and the state of preservation, the analysis by Lil­
lie (1998a; 2020) was only able to produce tenta­
tive sex / age determinations of M? aged 50—60 
for individual 4 and an indeterminate adult aged 
35—45 designation for individual 14, so at this 
stage in the research agenda there is little value 
in attempting to establish why the diet of these 
particular individuals differed when compared to 
all other individuals studied from the Epipalaeo­
lithic to Mesolithic periods.
It should also be noted that at Fat’ma Koba 
in Crimea, δ13C and δ15N values of –18.3 ‰ and 
10.5 ‰ respectively, for human remains recovered 
from Layer 4 (which is of either Murzak-Koba or 
Kukrek culture periodisation; Budd, Lillie 2020, 
p. 292), has also been interpreted as represent­
ing a diet wherein the majority of the dietary 
proteins are obtained from terrestrial resources. 
The δ13C of –18.3 ‰ has been suggested as indi­
cating the limited inclusion of some C4 resources 
at this location, due to the fact that at the time 
this individual would have been exploiting the 
environment (ca. 9000—7500 BP) Crimea was a 
peninsula in a freshwater context within an arid 
steppe environment (ibid. 2002, p. 292).
There are additional individuals of Mesolithic 
date from the cemeteries of Dereivka I (Individ­
ual 84 — discussed above) and Osipovka (Indi­
vidual 20b). As noted above, the values for the 
individual from Dereivka I are towards to lower 
end of the range identified, being closer to the 
terrestrial dominated baseline than the majority 
of the individuals analysed for the Mesolithic. At 
6361—5879 cal BC (OxA-6161; 7270 ± 110 BP) 
this individual is placed, chronologically, to­
Fig. 1. Human and faunal δ13C and δ15N values for the Epipalaeolithic and Mesolithic cemete-
ries in the study region (after Budd, Lillie 2020)
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Lillie, M. C., Budd, C. E. Diet Isotope Analysis and Related Studies in Prehistoric Ukraine: Fact, Fiction and Fantasy
wards the end of the Mesolithic period. Unfor­
tunately the individual from Osipovka produced 
isotope ratios that must be considered anomalous 
at this stage of the analysis (as they are well be­
low the entire dataset for this period to date) with 
values of 21.0 ‰ for δ13C and 5.7 ‰ for δ15N. Given 
that the material from this individual comprised 
fragmentary post cranial skeletal elements the 
possibility that this represents a fauna value as 
opposed to a human diet isotope value cannot be 
discounted with certainty, and without assess­
ment using a technique such as ZooMs (Zooar­
chaeology by Mass Spectrometry) to establish 
species identification (Buckley et al. 2009), this 
value is currently discounted from discussions of 
Mesolithic dietary pathways.
2.2. neolithic Subsistence. During the Neo­
lithic period there is a continued reliance on fresh­
water resources, to the point where these domi­
nate the diet at certain locations (fig. 2), although 
variability in consumption is again in evidence. 
The data have demonstrated that, at certain lo­
cations, there are individuals who have nitrogen 
ratios that are indicative of a reliance on terres­
trial resources, as opposed to the predominant di­
etary pathways wherein a reliance on freshwater 
resources is attested.
In this context, as can be seen from fig. 2, two 
individuals from the cemetery of Dereivka I have 
δ13C values of 23.4 ‰ and 21.7 ‰, and δ15N val­
ues of 9.9 ‰ and 10.5 ‰ respectively (individu­
als 49 and 33). In addition, individual 93 from 
Dereivka I has a similarly low nitrogen value at 
9.54 ‰ and a carbon value of 26.78 ‰, the latter 
value is very depleted but it is consistent with the 
faunal and fish values that have been obtained 
from this region. Other individuals at this loca­
tion have nitrogen values that, whilst towards 
the range of diets with a freshwater component 
in the Dnieper Rapids region in one instance, 
may well be more terrestrial based in light of the 
available baseline data for the forest-steppe zone 
(e. g. individuals 30 and 42). The values for these 
individuals are 23.61 ‰ and 23.2 ‰ for δ13C and 
12.07 ‰ and 11.08 ‰ for δ15N respectively.
There are two individuals from Vasilyevka V 
with δ15N values of 10.0 ‰ and 10.1 ‰, and a 
third individual who is elevated slightly at 10.6 ‰ 
(Individuals 8, 29 and 10), which would indicate 
the consumption of terrestrial based diets. Over­
all, it does appear that in general, δ15N values 
around ca. 11 ‰ and below are indicative of diets 
that were weighted more towards terrestrial than 
aquatic resources.
Of some considerable interest in terms of the 
dietary δ15N values that are recorded, are that 
two individuals from the cemetery of Yasinovatka 
(fig. 2) appear to have δ13C and δ15N that would ap­
pear to indicate divergent dietary pathways to the 
majority of this population (Budd et al. 2020).
It is worth noting that an additional individ­
ual 26, at Yasinovatka also produced a low δ15N 
Fig. 2. Human and faunal δ13C and δ15N values for the Neolithic cemeteries in the study region 
(after Budd, Lillie 2020)
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value at 7.6 ‰, but this individual was out of the 
acceptable range for C:N at 4.6, and as such, until 
further analysis (e. g. re-analysis for diet isotopes 
and ZooMs) can be undertaken, this individual is 
also discounted from further consideration.
Overall, whilst there is clear evidence to sup­
port variability in dietary pathways during the 
Neolithic (and preceding Mesolithic) period, it is 
apparent that that we need a considerable degree 
of caution in accepting that all samples analysed 
are in fact human, especially in the case of frag­
mentary material, or, as suggested by the current 
research, when the available fauna values indi­
cate that human nitrogen values below ca. 4—
5 ppm may suggest misidentification of fragmen­
tary skeletal materials.
2.3. eneolithic Subsistence. During the Ene­
olithic period the fact that a number of culture 
groups are available for study affords insights 
into differing food procurement strategies, whilst 
also highlighting the significant dietary differ­
ences that occur between groups following agro-
pastoral as opposed to mixed hunting, fishing and 
foraging food procurement pathways (fig. 3).
The data obtained from Igren 8 and Molukhov 
Bugor are clearly very distinct, isotopically, when 
compared to Verteba Cave (fig. 3). Of course, the 
fact that Verteba Cave is in western Ukraine, as 
opposed to the Dnieper region, will exert an influ­
ence on the isotope ratios in evidence, as will the 
fact that the individuals from Verteba are pre­
sumed to be Trypillia culture farmers. This cul­
ture, whilst having evidence to support the fact 
that varied subsistence regimes were practiced 
(e. g. Cuik 2008 and references therein), were 
essentially agro-pastoralists (see discussion be­
low).
The data from the Eneolithic phase at Igren VIII, 
which is dated to ca. 4337—4074 cal BC, were 
originally analysed by Rowena Henderson using 
high-resolution, incremental isotope analysis of 
the dentine in order to assess the relative con­
tribution of dietary proteins (Henderson 2015; 
Lillie et al. 2016). The δ13C and δ15N values from 
Igren VIII are the post-weaning diet signal at 
12 years of age for the ten individuals studied. 
These values, at 22.48 ± 0.9 ‰ and 14.02 ± 1.1 ‰ 
respectively, clearly indicate the significant con­
sumption of freshwater aquatic proteins, such 
as fish and molluscs (Budd, Lillie 2020). By con­
trast, the data from the chronologically later site 
of Molukhov Bugor, at ca. 3951—3705 cal BC, ex­
hibit carbon and nitrogen isotope values for the 
human inhabitants at δ13C = 21.8 ± 0.9 ‰ and 
δ15N = 12.2 ± 0.7 ‰, which, whilst indicative of 
the continued importance of aquatic proteins well 
into the Eneolithic period, are suggestive of more 
diversified subsistence strategies at Molukhov 
Bugor. As noted by Budd and Lillie (2020, p. 297), 
the more positive carbon, and lower nitrogen val­
ues at Molukhov Bugor probably reflect a mixed 
subsistence base, where aquatic proteins were 
Fig. 3. Human and faunal δ13C and δ15N values for the Eneolithic cemeteries in the study region 
(after Budd, Lillie 2020)
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Lillie, M. C., Budd, C. E. Diet Isotope Analysis and Related Studies in Prehistoric Ukraine: Fact, Fiction and Fantasy
consumed alongside wild and domestic herbiv­
ores, or that there is possibly a difference in the 
composition of fish species available at the Sula 
river. By contrast, the data from Igren VIII are 
indicative of the heavy consumption of freshwa­
ter proteins with no isotopic or material culture 
evidence to suggest the consumption of dietary 
proteins from the terrestrial landscape in any sig­
nificant quantity (Telegin, Zaliznyak 1975; Hend­
erson 2015; Lillie et al. 2016; 2020).
3. Radiocarbon dating and the absolute 
chronology of the Dnieper-Donets Mar-
iupol-type cemeteries. It is an established 
fact that there is a freshwater radiocarbon res­
ervoir effect (FRE) influencing 14C dating of the 
Dnieper-Donets Mariupol-type cemeteries (Lillie 
et al. 2009). However, despite the fact that there 
is clear evidence for a freshwater radiocarbon 
reservoir effect in the Dnieper River region, it is 
naпve to suggest that it is a simple case of calcu­
lating a «linear correction factor» using the exist­
ing limited dataset alongside δ15N values to ac­
count for the 14C offset for these cemeteries (e. g. 
Kotova 2018).
The freshwater reservoir effect, in oversimpli­
fied terms, can cause an anomalous 14C offset in 
the radiocarbon age of a sample that is either 
from a freshwater context, or is from a human / 
animal that has consumed freshwater resources; 
specifically the effect can produce a 14C age that is 
artificially «older» than its true radiocarbon age 
(Ascough et al. 2005; Philippsen 2013).
Radiocarbon dating at three sites in the 
Dnieper region has revealed a 14C reservoir ef­
fect of ca. 100—500 years (see Lillie et al. 2009, 
p. 62, table 4). At Yasinovatka, there is a chron­
ological offset between the fish tooth sample at 
6840 ± 37 uncal BP (OxA-17498), and the deer 
tooth sample at 6121 ± 34 uncal BP (OxA-17500), 
of about 720 years (Lillie et al. 2009, p. 261). 
This equates to a ca. 250 uncal year difference 
between the fish sample and the human sam­
ple which is dated to 6593 ± 35 uncal BP (OxA-
17499), and ca. 470 years between the human 
sample (with the latter having a ratio of 22.6 ‰ 
for δ13C and 14 ‰ for δ15N), and the deer sample. 
By contrast, at Dereivka I, the chronological dif­
ference between the fish sample (burial 29) at 
6915 ± 50 uncal BP (OxA-17501) and the deer 
sample (square 801, burial 29) at 6147 ± 35 uncal 
BP (OxA-17594) is approximately 770 years. This 
equates to a disparity of 517 years between the 
fish and human sample which is dated to 6398 ± 
35 uncal BP (OxA-17495), and 251 years between 
the human and the deer sample. The human sam­
ple (individual 29) from Dereivka I has yet to be 
analysed for diet isotopes, but the range from 12 
individuals at this cemetery for δ15N is 9.9 ‰ to 
13.05 ‰ (δ15N 11.96 ± 0.95 ‰).
Given that the Yasinovatka individual has a 
δ15N value of 14 ‰ and an offset from the terres­
trial baseline (fauna) of 470 years, we must con­
sider that, in theory, at Dereivka I the FRE offset 
could be anything from zero (9.9 ‰ being a ter­
restrial diet signature) through to a ca. 350 year 
offset based on the human values obtained to 
date. However, it is important to emphasise that 
we cannot assume the degree of 14C offset is sys­
tematic across the Dnieper region, or assume re­
source equivalence in terms of δ15N baseline val­
ues between Yasinovatka and Dereivka I, which 
are located in different ecozones and different 
parts of the Dnieper system.
There are a number of challenges associated 
with unravelling freshwater radiocarbon reser­
voir effects at prehistoric sites (see Guiry 2019 
for a review of the biogeochemical processes in­
volved). Furthermore, there is an open debate 
in the academic literature concerning which 
statistical technique to apply to calculate 14C off­
set. Most commonly, various forms of linear re­
gression models are used (Schulting et al. 2014; 
2015), with other studies preferring a Bayesian 
approach (Bronk Ramsey et al. 2014; Sayle et al. 
2016).
A dedicated study investigating the freshwa­
ter radiocarbon reservoir effect at a number of 
prehistoric micro-regions in Lake Baikal, Siberia 
has shown that developing a correction for the 
FRE is not a straight-forward process. Weber et 
al. (2016) analysed 42 pairs of radiocarbon ages 
from human bone and terrestrial herbivore bone 
or tooth samples alongside δ15N and δ13C values 
to calculate a correction factor using the linear 
regression methods developed by Schulting et al. 
(2014; 2015). Even with the significantly larger 
dataset, and multiple technique approach, they 
note that not only does the predictive power of 
these regression models vary notably between 
micro-regions (in some cases the regression equa­
tions explain less than 50 % of the variation ob­
served), there are also a number of cases where 
δ13C values are better predictors of 14C variation 
than δ15N values (Weber et al. 2016, p. 82).
To reiterate, the comprehensive freshwater res­
ervoir effect study at Lake Baikal has 42 paired 
dates within an overall dataset of 256 AMS de­
terminations, and despite this dataset the au­
thors could not fully resolve the 14C offset. In the 
Dnieper region we have 2 paired dates. As such, 
it is apparent that, at present, there is no simple 
correction factor that can be uniformly applied to 
radiocarbon ages across the Dnieper River cem­
eteries.
4. Verteba Cave. Recent research undertaken 
at the site of Verteba Cave in western Ukraine 
has resulted in the production of a number of pa­
pers, by different research teams, following on 
from the initial investigations by Alex Nikitin 
and Mykhailo Sokhatsky between 2005 and 2008 
(Nikitin et al. 2010; 2017; Nikitin 2011). Whilst 
there is no denying the fact that this location is 
exceptional for many reasons, not least due to the 
extremely long use of this cave as a location for 
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the deposition of human remains, certain aspects 
of the research that has been produced from in­
vestigations at this location require redress. 
Amongst the topics that have arisen from the 
recent research at this site, and which could be 
seen to require some further consideration, is the 
suggestion that Verteba Cave can be used to as­
sess the shift from fisher-hunter-forager subsist­
ence economies to farming economies in Ukraine. 
In this context Karsten and co-workers (e. g. 
Karsten et al. 2014; 2015b) have contrasted the 
pathologies in evidence at Verteba to those iden­
tified by Lillie (1998a) in the Dnieper region, sug­
gesting that these pathologies demonstrate the 
impact of the transition to farming in Ukraine on 
overall health status.
The fact that there are consequences in terms 
of oral and overall health status at the transi­
tion to farming is a well-established fact, but, the 
point is that the pathologies that are reported for 
the individuals studied at Verteba, which are all 
assumed to be Trypillian farmers (an observation 
that is brought into question below), do not rep­
resent a continuum, in subsistence terms, for the 
populations of Ukraine as they transition from 
hunting, fishing and foraging to farming. One 
key issue lies in the fact that Verteba is ca. 500—
600 km west of the Dnieper Rapids region, where 
Lillie (1998a) undertook his study, and a second 
issue is that Trypillia groups are not the popula­
tions who develop from the groups interred in the 
Dnieper-Donets Mariupol-type (DD M-t) cemeter­
ies, which is convincingly proved by anthropologi­
cal and archaeogenetic studies (Potekhina 2018). 
The hunter-fisher-forager groups in and around 
the Dnieper do not transition to agriculture until 
much later, and even when we see forager-farmer 
interactions, only limited elements of the farm­
ing «package» are integrated into subsistence 
strategies. It is invalid to suggest that patholo­
gies recorded on the possible Trypillia (or chrono­
logically equivalent) individuals from Verteba (or 
those from any period between the Mesolithic and 
later historical periods), can be considered to be 
suitable for comparative purposes for the simple 
fact that the subsistence differences and socio-
cultural trajectories of the DD groups interred 
in the Mt cemeteries are distinct from Trypillia, 
and cannot realistically be compared because DD 
groups do not develop farming; and ultimately, 
partially due to a failure to adapt, Trypillia fails 
(Kruts 2012, p. 230).
Referring back to the suggestion that it is er­
roneous to assume that all of the individuals 
studied at Verteba are Trypillian in origin, we 
must return to the issues surrounding the lack 
of absolute dating of the remains from Verteba 
as discussed by Lillie et al. (2015; 2017; 2020), 
and whilst Madden et al. (2018, p. 901) have pre­
viously stated the following: «Lillie et al. (2017) 
have suggested dating and isotopic testing for all 
remains at the cave to ensure their provenance, 
while this would improve our understanding of 
the use of the cave and the Tripolie (Trypillia) cul-
ture it would also require destruction of what is a 
unique sample. This population is a non-renew-
able resource informing us of the past, we must 
be careful to balance potential knowledge gained 
with current technology against preservation of 
the remains for the future (Makarewicz and Sealy, 
2015)».
This is a somewhat trite statement which be­
lies the fact that the approaches adopted by these 
researchers at Verteba have failed to adequately 
demonstrate that the remains being studied are 
contemporary, and as Ledogar et al. (2019) clear­
ly demonstrate, Mesolithic through to modern 
cultural material and human remains are in evi­
dence at this location. In actual fact, and appar­
ently contra Madden et al. (2018, p. 901), Ledogar 
et al. (2019, p. 141) state that understanding the 
chronology of Verteba cave is integral to further 
interpretations of the material evidence from 
Verteba (our emphasis). The simple fact is that 
all of the studies that have been produced that 
group together undated material from secondary 
contexts are fundamentally and methodologically 
flawed, and these studies have simply failed to 
meet the basic requirements of rigour in scientific 
approach that should be anticipated.
Logic and good science dictates that when the 
available evidence clearly indicates that strati­
graphical integrity cannot be assured and that 
co-mingling of sequences has been clearly dem­
onstrated (as outlined in detail below), treating 
the material from Verteba as an homogenous col­
lection of Trypillia individuals for the purposes 
of producing a seemingly numerically significant/
satisfactory dataset is unjustifiable, at best. The 
fact that the studies prior to Ledogar et al. (2019) 
perpetuate these unsound assertions is perhaps 
made more problematic given that a number of 
other studies (e. g. Lillie et al. 2015; 2017) em­
phasised that, in light of the evidence from these 
studies and those of Nikitin (2011; Nikitin et al. 
2010; 2017) absolute dating is essential to ensure 
that the conclusions of previous studies (e. g. 
Karsten et al. 2014; 2015a; 2015b) into the palae­
opathological markers in evidence are accurate, 
and that all of the individuals used in these stud­
ies are, in fact, Trypillian in date.
Further issues arise in the reporting of the 
material from Verteba Cave with the observa­
tion that in their 2019 study Ledogar et al. (2019, 
p. 151) state that «Probability distributions for 
the 14C dates suggest that there may have been 
either two periods of occupation, or two discrete 
events resulting in deposits during the Eneolithic» 
(fig. 4). As such, with this basic observation in 
place, it is clear that the grouping and treating 
of the Verteba material as an homogenous as­
semblage immediately fails to account for the fact 
that the pathologies in evidence actually repre­
sent at least two phases of deposition.
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Lillie, M. C., Budd, C. E. Diet Isotope Analysis and Related Studies in Prehistoric Ukraine: Fact, Fiction and Fantasy
However, of even greater concern is the fact 
that Lillie et al. (2017, p. 313) had previously stat­
ed that their «Figure 4 highlights the fact that the 
calibrated dates separate out into what appears to 
be three discrete phases of activity, an observation 
that would be consistent with the evidence from 
the palaeopathology» (evidence for healed trauma 
suggesting that low levels of repeated interper­
sonal violence are attested at Verteba; fig. 5; 6). 
Of note here is the fact that no mention is made 
of the previous studies undertaken by Lillie and 
co-workers (2015; 2017) by Ledogar et al. (2019), 
who instead chose to focus on the stable isotope 
work by Lillie and co-workers, as this does not 
undermine the premise of their 2019 paper. In ad­
dition the AMS dates in Lillie et al. (2015; 2017) 
also extend the period of use at Verteba, back 
to pre 3900 cal BC, thus reinforcing the work of 
Nikitin et al. (2010). Importantly, as the Trypil­
lia material is co-mingled with material of both 
earlier and later date, and there are at least two, 
and possibly three depositional events at Verteba 
during the Trypillia phases, it is an established 
fact that stratigraphic integrity simply cannot 
(and should not) be assumed for the material cul­
ture and human remains from Verteba.
As noted by Lillie et al. (2020), a total of 33 
conventional dates from the Kiev radiocarbon fa­
cility, were reported in Nikitin et al. (2010), 14 
of which were made on human remains (bone 
collagen) from the cave site. These dates prima­
rily spanned the periods ca. 3600—2500 cal BC 
(Trypillia CI—CII), but large errors, e. g. Ki14308 
at 4910 ± 400 BP (rib fragment), extended the 
range to 4652—2674 cal BC. The combined dates 
on pottery and human bone did extend the use 
of the cave back into the period of activity of the 
Shypynetska culture grouping of Trypillia, who 
used the cave ca. 3745—3550 cal BC (i. e. prima­
rily into Trypillia stage BII). Fundamentally, the 
work of Nikitin et al. (2010) and Lillie et al. (2017) 
has clearly demonstrated that the earlier dates for 
Trypillia activity equate to stage BII of Trypillia 
and that subsequent activity extends the periods 
of cave use into stage CI and CII of this culture. 
Lillie et al. (2020) note that in addition to refining 
the earlier work of Karsten and co-workers this 
is also clearly contra Kadrow and Pokutta (2016, 
p. 3) who state that «the oldest traces of human 
activity at the Verteba Cave come from the late 
CI phase and are associated with the Shipentsy 
group, living near the Badrazhy group on the cen-
tral Dniester Plateau». Importantly, Nikitin et al. 
(2010, p. 15) also noted that the use of the cave 
was not continuous, as attested to by sterile hori­
zons between cultural layers, and that use by the 
Fig. 4. Probability distributions for the calibrated 
dates for the eight samples from Verteba Cave that 
are associated with the Eneolithic (after Ledogar et al. 
2019)
Fig. 5. AMS distribution plots for the individuals analysed by Lillie et al. (2015; 2017)
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Koshylovetska and Kasperivska groups occurred 
between ca. 3550—2747 cal BC.
In addition to the evidence from the radiocarbon 
dating at Verteba, DNA analysis undertaken by 
Nikitin and co-workers (e. g. Nikitin et al. 2017) 
has shown that the individual’s analysed exhibit a 
typical Neolithic farmer package of mitochondrial 
DNA (mtDNA), with lineages traced to Anatolian 
farmers and Neolithic farming groups of central 
Europe. At the level of mtDNA haplogroup fre­
quencies the TC population from Verteba demon­
strates a close genetic relationship with popula­
tion groups of the Funnel Beaker / Trichterbecker 
cultural complex from central and northern Eu­
rope (ca. 3950 ± 2500 BCE). Furthermore, two 
specimens belong to haplogroup U8b1 at Verteba, 
and these individuals can be viewed as a connec­
tion of Trypillia Culture (TC) groups with the Up­
per Palaeolithic populations of Europe.
The DNA analysis obviously provides impor­
tant insights into the genetic origins of the Trypil­
lia groups that used Verteba, but, importantly, 
an additional issue is highlighted by Nikitin’s 
work. This relates to the suggestion that the dat­
ing and stable isotope values obtained by Ledogar 
et al. (2019) on a 2nd and 4th metacarpal, are from 
discrete individuals, an observation that is sub­
ject to question. DNA analysis by Nikitin (pers 
comm. 16/11/2019) has already shown that there 
are two metacarpals (left and right hand) exca­
vated in May 2008 which appear, genetically, to 
be from the same individual. Importantly for the 
current discussion is the observation that one of 
the metacarpals was found near the surface (al­
luvial deposit, level 1) in the SE corner of quad­
rant 1 at Verteba, whilst the other was found at 
level 3 (within or on top of the clay matrix of the 
cave floor) in the NW corner. As both elements 
appear (genetically) to belong to the same person, 
the obvious conclusion must be that there is some 
considerable mixing between levels. Given the 
comingled, secondary depositional, context of the 
material at Verteba, caution is clearly warranted 
when analyzing material that cannot be shown to 
be from discrete individuals.
Thus, the observation that the dating and sta­
ble isotope values obtained on a 2nd and 4th met­
acarpal, are from discrete individuals at Verteba 
Cave (Ledogar et al. 2019, p. 148, table 2, fig. 3) 
should be queried (see table 1), especially given 
the close correspondence of the isotope values 
from these «individuals» (δ13C of –19.8 ‰ and 
19.8 ‰ and δ15N of 9.8 ‰ and 9.9 ‰ respectively), 
and the identical AMS dates obtained.
Importantly, Ledogar et al. (2019, p. 152) 
state that «The 14C ages for bone samples from 
site 20, 4955 ± 25 BP (CC167135), 4965 ± 25 BP 
(CC167136), and 4900 ± 20 BP (CC169195), 
were compared and are not statistically different 
(T = 0.78, χ2 = 5.99 at α = 0.05; df = 2) suggesting 
that the strata from these deposits are contempo-
raneous». The problem appears to be that there is 
an identical date being reported from a different 
context. However, the subsequent statement by 
these authors (2019, p. 152) that CC169195 is in 
fact from site 20 and not location Г3 is problemat­
ic, and perhaps highlights some confusion in the 
reporting of these finds (which can happen obvi­
ously). As noted above, the isotope data and AMS 
dates would strongly suggest that these elements 
are from the same individual. Consequently, and 
given the lack of stratigraphic integrity at Verte­
Fig. 6. Calibrated dates plotted on the 
radiocarbon curve — note that the loca­
tion of the dates would indicate at least 
two or three discrete phases of deposition 
for these skulls (from Lillie et al. 2017)
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ba, a note of caution is warranted both in terms of 
methodological approaches that assume that all 
elements are from discrete individuals (a clearly 
untenable assumption), and where near identical 
data is generated the possibility that the same in­
dividual has been analyzed must surely be taken 
into account (a point that is touched upon further 
below).
As the evidence indicates, the remains at 
Verteba are secondary deposits, as such, the 
finds of smaller elements from the same individ­
ual suggests that the remains were relocated to 
Verteba prior to full disarticulation occurring at 
the primary point of interment / processing. This 
actually adds another interesting new aspect to 
the rituals in evidence at Verteba, and would cor­
roborate the observations by Lillie et al. (2015; 
2017) that the processing and inclusion of the 
remains into the cave deposits probably occurred 
relatively soon after the death of the individual.
With the above observations in mind it is ob­
viously equally fictitious to assert that all of the 
Table 1. Identical AMS dates for elements from differing locations at Verteba Cave (after Ledogar et al. 2019)
Site L/F Lab # Sample # Taxon Element Age Fraction Modern
14C, BP 2σ date, cal BC
20 L2 CC169199 C15-VC13 Homo sapiens 4th metacarpal A 0.54 4900 ± 20 3704—3648
Г3 L3 CC169195 C16-VC12 Homo sapiens 2nd metacarpal A 0.54 4900 ± 20 3704—3648
Abbreviations, here and in table 4: L — level, F — feature, Ob — overburden, Sf — surface, A — adult, S — 
subadult.
Table 4. Identical AMS dates for elements from differing locations at Verteba Cave (after Ledogar et al. 2019)
Site L/F Lab # Sample # Taxon Element Age Fraction Modern
14C BP 2σ date cal BC/AD
Г3 Sf. CC-167139 C15-VC09 Homo sapiens Parietal A 0.85 1315 ± 25 656—780, 76 %; 
741—766, 24 %
cal AD
Г3 L.1 CC-169196 C16-VC17 Canis familiaris Metapodial A 0.73 2505 ± 20 778—730, 22 %; 
692—659, 17 %; 
651—543, 61 %
cal BC
Table 2. Strontium, oxygen and carbon isotopic composition of tooth enamel of three individuals  
from Verteba Cave (after Madden et al. 2018)
Sample Element Age, years Tooth formation, crown complete, years Sex 87Sr/86Sr
δ13CvPDB, 
‰
δ18OvSMOW, 
‰
EA14 3rd left maxillary molar 17—25 12—16 Male 0.70975 11.8 23.8
EA15 Canine 25—35 6—7 Male 0.70931 11.9 23.4
EA16 Left lateral maxillary incisor 25—35 4—5 Male 0.70966 11.5 23.7
Table 3. Sr, δ13C and δ18O results from human and pig samples at Verteba (after Lillie et al. 2017)
Species Individual Sample ID Element Sex 87Sr/86Sr δ13C δ18O
Human 7 V1.1.1 Max right M2 M 0.709660 13.12 5.80
Human 3 V3.18.1 (1.2) Max left P3 F 0.709741 12.86 6.98
Human 4 V3.14.1 Max right P4 M 0.709616 11.99 6.83
Human 6 V3.15.1 Max Right M1 M 0.709692 12.86 6.48
Human 2 V3.16.1 Max right P3 M 0.709312 12.64 6.31
Human 1 V3.17.1 Max Left M1 M 0.709842 12.30 6.39
Human 8 A22 Incisor I 0.709344 12.17 5.83
Human 9 M5 Mand right PM2 M 0.709475 9.12 5.94
Pig-F2 — F2.6.2 Incisor — 0.709100 — —
Pig — F2.6.4 Mand right PM2 — 0.709700 — —
Text Highlighted in Grey are samples that have identical 87Sr/86Sr ratios to samples EA16 and EA15 in the 
Madden et al. (2018) study: discussed in the text
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Антропологія
trauma in evidence at Verteba can be wholly as­
signed to a single stage of the Trypillia culture, as 
the available dating contradicts this, and equally, 
the lack of absolute dating of comingled / mixed 
deposits precludes such an assertion. It is appar­
ent that the failure by Karsten and co-workers 
to date every sample (prior to the Ledogar et al. 
2019 paper) suggests that there was a working 
assumption that the remains were Trypillian in 
age and as such, contemporary, but this is dis­
proven. Additionally, there is a knock-on effect 
to all previous studies by these authors in that 
the grouping of the material from Verteba, and 
the implied contemporaneity, is at best an error 
of judgement, and at worst misleading. Similarly, 
suggesting that the available evidence attests 
to a CII age for the trauma in evidence is also 
somewhat economical with the facts, in that the 
dating by Nikitin (2011; Nikitin et al. 2010, and 
subsequently Lillie et al. 2015; 2017) had already 
shown / has shown that the material that equates 
to Trypillia is also dated to stages BII and CI of 
this culture (and possibly slightly earlier), thus 
establishing the fact that inter-personal violence 
occurs before the decline of Trypillia. This fact is 
actually far more interesting in research terms, 
as the available evidence indicates that low level 
endemic violence may characterise this farming 
culture across the main stages of its evolution, an 
observation that was not possible prior to the dis­
covery of Verteba Cave, as the skeletal record for 
Trypillia is lacking in general, and very poor for 
the earlier stages of this culture.
Further irregularities are highlighted between 
the studies of Lillie et al. 2017 and Madden et 
al. 2018 in the reporting of strontium isotopes 
from Verteba. In their 2018 study Madden et al. 
(2018, p. 899) present the data in table 2 showing 
the strontium, oxygen and carbon isotope results 
from three individuals at Verteba.
Previously Lillie et al. (2017, p. 316) produced 
the results in table 3 for 8 human for 8 human 
and 2 pig samples from Verteba Cave.
Whilst the δ13C and δ18O ratios differ for two of 
the individuals analysed by Madden et al. (2018) 
it is intriguing to note the identical 87Sr/86Sr ra­
tios between individual EA15 in the Madden et 
al. (2018) study and individual 2 in the Lillie et 
al. (2017) study, and the identical ratio for indi­
vidual EA16 when compared to individual 7 in 
the Lillie et al. (2017) study (table 3: highlight­
ed text). Whilst the δ13C and δ18O ratios are not 
comparable between these two studies, it might 
be worthwhile identifying whether the individu­
als with identical strontium values are in fact the 
same individuals, as given the finite nature of the 
resource, it is clear that different research teams 
need to work in synergy in order to reduce repli­
cation of analyses and thus propagate a reduction 
in the available resource. Of course, the fact that 
two individuals have produced identical Sr ratios 
is also somewhat unusual in itself.
5. Additional Observations. Ledogar et al. 
(2019, p. 149) observe that at Verteba «Nitrogen 
values for two of the humans, two dogs, and one 
pig are at the same trophic level (9.0—10.5 ‰) as 
the freshwater species, while the other humans, 
pig, dog, and chicken (2.7—8.4 ‰) fall below the 
trophic level of the freshwater species». The dat­
ing, alongside the nitrogen diet isotope values 
of 2.7 ‰ and 2.9 ‰ for human and dog remains 
from Verteba suggest that the Г3 site represents 
a mixed location with dates of the dog across three 
periods between 778—543 cal BC and the human 
placed across two periods between 656—780 cal 
AD (Iron Age and Scythian; table 4).
Whilst at face value the 2.7 ‰ and 2.9 ‰ ni­
trogen isotope values for the Iron Age dog and 
Scythian human appear to be reliable results, 
the fact that Ledogar et al. (2019) note that they 
are inconsistent with the data from Murphy et al. 
(2013), and in fact are inconsistent with all other 
studies undertaken on the remains from Ukraine 
to date, is potentially cause for concern.
It is clear that the application of a technique 
such as ZooMS (Zooarchaeology by Mass Spec­
trometry) should be considered to verify the iden­
tification of these samples. The isotope values are 
in fact well below all those of previous studies 
and also generally well below those in Ledogar 
et al. (2019). As such, stating that they suggest a 
«strongly herbivorous diet» (2019, p. 154) needs 
to be tested as these values do not reflect troph­
ic enrichment (which would be expected) or the 
nature of the data that has been generated up 
to this point. In general, whilst the majority of 
the data at Verteba is consistent with the ratios 
that would be expected for humans consuming 
terrestrial diets wherein C3 resources are being 
consumed, and the data are consistent with the 
levels identified by Budd and Lillie (2020) for the 
Mesolithic through to Eneolithic periods further 
east in the Dnieper Basin, that are associated 
with terrestrial diets, the values of 2.7 ‰ and 
2.9 ‰ need to be investigated further given their 
anomalous nature in relation to the available 
data from Ukraine as a whole.
One further observation is that Ledogar et al. 
(2019, p. 153) state that «Populations in Meso-
lithic Ukraine have been considered hunter-fish-
er-gatherers who relied heavily on animal pro-
tein; however, the isotopic data from individual 
CC167136 from Verteba Cave suggests that he or 
she consumed much more plant protein. While 
the data from this study is only the diet of one 
individual, it does introduce the possibility that 
diet was quite variable across the region at this 
period». Lillie et al. (2009; 2010; Lillie 1998 and 
a number of other papers by these authors) had 
actually already stressed that Mesolithic diets did 
not rely heavily on animal protein, and they had 
also already noted the fact that variability in diet 
was occurring, and in actuality the data continues 
to highlight this (Budd, Lillie 2020). The reader is 
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directed towards Lillie et al. (2020), Lillie (2020) 
and Budd and Lillie (2020) for a detailed critique 
of the Ukrainian evidence.
6. Discussion. The identification of an em­
phasis on aquatic resources from the Epipalaeo­
lithic period onwards in the Dnieper region is 
significant for Ukraine as it is generally assumed 
that broad-based resource procurement strate­
gies are more characteristic of later, post-glacial 
or Mesolithic groups (Dolukhanov 2000, p. 78). 
Importantly, as noted by Lillie (2003a), it also 
suggests that the preoccupation with the exploi­
tation of large game animals, that characterises 
discussions of prehistoric subsistence strategies 
in Ukraine (Balakin, Nuzhinyi 1995; O’Connell et 
al. 2000), fails to account for the reality, especially 
given the considerable evidence for the exploita­
tion of aquatic resources; resources which clearly 
formed an essential element of Epipalaeolithic 
through to Eneolithic diets across the early to 
mid-Holocene. It is also likely that fish-based pro­
teins periodically increased in significance across 
the Epipalaeolithic to Eneolithic periods for vari­
ous socio-economic, environmental and even ritu­
alised reasons. To reiterate however, it should be 
remembered that the designation of «Neolithic» 
for Ukraine does not necessarily equate to the 
«traditional» observation of an associated shift 
towards the exploitation of domesticated plants 
and animals, and as such continuity in subsist­
ence practices across the early to mid-Holocene is 
not atypical for many regions of Eastern Europe.
The fact is that fish, and freshwater resources, 
are an important dietary staple from the Epi­
palaeolithic period onwards, and it would be fic­
titious to suggest that the exploitation of these 
resources represents the exploitation of a «star­
vation food» that is only relied upon during pe­
riods of resource stress. Amongst the fantasies 
that are highlighted in dietary studies is the idea 
that meat consumption is the «be all and end all» 
of the subsistence spectrum during the earlier 
Holocene. Furthermore, it appears that we see 
diversification in subsistence strategies between 
the Epipalaeolithic and later Mesolithic periods 
as the data suggest that a broader dietary spec­
trum was being exploited in the latter period as 
plant and animal resources expand and diversify 
(Budd, Lillie 2020). The isotope data also dem­
onstrate that as early as ca. 7000—6600 cal BC 
there is evidence to suggest that certain individu­
als, in this case at the cemetery of Marievka, were 
consuming diets that were predominantly based 
on the consumption of terrestrial as opposed to 
freshwater resources (ibid. 2020). Throughout 
the Neolithic period the data again indicate that 
a heavy reliance is placed upon the exploitation 
of freshwater resources, especially at the sites 
of Yasinovatka, Nikolskoye and Dereivka I. At 
Yasinovatka, Vasilyevka V, and Dereivka I we 
again have individuals who were consuming ter­
restrial as opposed to aquatic dominated diets. 
By the Eneolithic period diversification is occur­
ring. The isotope analysis at Igren VIII indicates 
a heavy reliance on freshwater resources, but at 
Molukhov Bugor the data indicate that more var­
ied diets were being consumed, albeit with fresh­
water resources still forming an integral element 
of subsistence strategies. The main point of de­
parture, as might be anticipated, is the evidence 
from Verteba Cave in western Ukraine, where 
the diet isotope data supports the exploitation of 
terrestrial dietary pathways for the (presumed) 
Trypillia farmer groups who interred their dead 
in secondary contexts at this location.
As has been emphasised elsewhere (Lillie et al. 
2009), the heavy reliance on freshwater resources 
across the Epipalaeolithic through to Eneolithic 
periods in and around the Dnieper river system 
in Ukraine is resulting in a radiocarbon fresh­
water reservoir effect (FRE) that is impacting on 
the dating of the individuals interred in the DR 
M-t cemeteries of Ukraine. The discussion above 
has highlighted the fact that there is no simple 
correction factor that can be applied at the whole 
cemetery level due to the variations in δ15N val­
ues that are in evidence. However, the fact that 
there are individuals at a number of cemeteries 
who consumed diets that were dominated by ter­
restrial resources, and who exhibit values for δ15N 
across the range of ca. 9—11 ‰, does offer some 
potential for future studies aimed at refining the 
FRE correction factors at certain locations.
Finally, the discussion throughout this paper 
has sought to highlight areas where research 
agenda’s overlap, and where the research itself is 
occasionally perhaps lacking somewhat in meth­
odological and scientific rigour. In this context the 
research undertaken at Verteba Cave has proven 
fruitful in terms of a critique of methods and the 
dissemination of results. Whilst this overview is 
envisioned as an appraisal of the state of play up 
to this point, it is not intended to be critical per 
se, as there are many pitfalls in the development 
of new and innovative research agenda’s. This is 
especially so in regions such as Ukraine, where 
the history of research makes it difficult to gain 
a rapid overview of cultural developments, and 
where the application of scientific methodologies 
such as AMS dating and isotope analyses are still 
relatively new in studies of past human groups. 
The costs involved in the application of these 
techniques can be prohibitive, and it is clear from 
the studies at sites like Verteba Cave in west­
ern Ukraine, that the work of different research 
teams can intersect, and that this can on occa­
sion lead to inconsistencies and overlaps in the 
reporting of research findings. Undoubtedly con­
certed research efforts aimed at ensuring synergy 
in approach and securing funding for long-term 
studies must form an integral aspect of future re­
search in Ukraine.
Conclusions. At present the majority of the 
cemeteries studied from the Dnieper Basin ex­
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Антропологія
hibit either long periods of interment, or multiple 
phases of use, or both, this is the case for many 
DD/M-t cemeteries, and is also true for Verteba 
Cave in western Ukraine. Verteba Cave itself is 
unique in the myriad ritual activities that are 
attested, and in offering insights into the Trypil­
lia culture that would have been impossible to 
disentangle without such a location. Of some 
significance is the observation that low levels of 
endemic inter-personal violence may well have 
characterised this population across stages BII to 
CII of its existence, and of course it is conceivable 
that such activities may have characterised the 
Trypillia culture in general.
The data has shown that in the Dnieper region 
the exploitation of freshwater resources is occur­
ring from the Epipalaeolithic through to Eneolithic 
periods, and whilst there is some variability in evi­
dence, prehistoric diets were not simply focussed 
on the exploitation of large game animals. Fresh­
water resources were stable resources in a chang­
ing environment and it is likely that all of the re­
sources of the riparian zone were being exploited 
across the Epipalaeolithic through to Eneolithic 
periods and beyond. Logically we now need to un­
dertake full cemetery analyses for all of the periods 
that have been studied to date, and to extend the 
analyses into the later prehistoric periods to try to 
elucidate the nature of the adoption of domesticat­
ed resources by subsequent culture groups, whilst 
also expanding the analyses further, beyond the 
areas that are currently being studied.
It is apparent that concerted collaborative ef­
forts are needed to ensure that the study of pre­
historic populations are systematic, integrated 
and targeted to ensure that realistic interpreta­
tions and conclusions are being generated from 
studies in the territory of Ukraine. It is also read­
ily apparent that replication of data is wasteful in 
terms of research time and resources, and given 
that Madden et al. (2018, p. 901) state that Verte­
ba is a «unique» and a «non-renewable resource» 
the potential analysis of the same individuals 
as identified above would reinforce the need for 
dialogue and synergy in approaches to such data. 
Similarly, the avoidance of misrepresentation 
or misuse of data from previous studies, which 
undermines the discipline and the peer review 
process somewhat, and weakens the value of the 
scientific research that has been undertaken over 
the past three decades (particularly so in the case 
of dating and isotope studies), is fundamental if 
the research being produced is to remain valid, 
authorised and significant.
Unfortunately, the fact is that despite our 
studies since 1992—1993, there is still a lot of 
analysis that needs to be undertaken if we are 
to adequately characterise the socio-cultural and 
economic development of the prehistoric popula­
tions of Ukraine, not least because the majority 
of our research has focussed on the populations 
in and around the Dnieper river system due to 
the exceptional preservation of human skeletal 
remains in cemeteries from the earliest Holocene 
through to ca. 3500 cal BC (and beyond) in this 
region. Throughout our research program a lack 
of resources has inhibited the degree to which 
dating and dietary isotope studies have been able 
to be applied. Ukraine has been shown to be ex­
ceptional in terms of its prehistoric cultural de­
velopments, the existence of cemeteries from as 
early as ca. 10,200 cal BC and their use across 
all periods of prehistory, the fact that the genetic 
data indicates that the Yamnaya culture is piv­
otal in terms of the origins of the Indo-Europe­
ans, and that it is likely that earlier Sredni-Stog 
culture groups may well be central to these de­
velopments (e. g. Mathieson et al. 2018). Future 
research agendas now need to expand upon the 
foundations that have been established and un­
dertake whole cemetery multi-disciplinary analy­
ses in order to further enhance our understand­
ing of socio-economic and societal developments 
during the early to mid-Holocene in Ukraine.
Acknowledgements. We would particularly 
like to thank Professor, Dr. Hab. Dmytro Telegin 
(to whom this paper is dedicated) and Dr. Inna 
Potekhina, as, without their help, guidance, col­
laborative spirit, inspiration, and advice the early 
research would have been far more difficult and 
onerous. We would also like to thank Alexei Ni­
kitin for discussions around the DNA analyses, 
and for advice and opinions in relation to our 
thoughts regarding Verteba cave. Also, Mykhailo 
P. Sokohatskyi was kind enough to invite MCL to 
visit Verteba in 2011, with Dr. Inna Potekhina, 
and Prof. Aleksandr Potekhin drove us from Kiev 
to Verteba, no easy undertaking given the road 
conditions away from the major urban centers.
Professor Lillie would also like to make men­
tion of two colleagues who are also no longer with 
us, but who both in their own ways helped him 
throughout the earlier years of his research — 
firstly Professor Timofeev (St. Petersburg) who 
helped with access to the collections housed in St. 
Petersburg and with an extended visit during the 
early part of the research agenda, and secondly 
Professor Dolukhanov, who was always helpful 
in offering advice and access to papers across a 
significant part of the research. Pivotal, in terms 
of Lillie’s academic development was, of course, 
his PhD supervisor the late Professor Marek Zve­
lebil, one of the more influential thinkers in ear­
lier Prehistoric studies. They are all missed.
Funding. A NERC-AHRC National Radio­
carbon Facility (NRCF) grant (to MCL) provided 
funding for the AMS Dating presented in the 
Lillie et al. 2015; 2017 papers (Project — No. 
NF/2011/2/18). In addition, WAERC (Wetland 
Archaeology & Environments Research Centre) 
at the University of Hull (now based at Umeå 
University in Sweden) funded a considerable 
amount of the dating and isotope work that has 
been undertaken since 1991—1992.
265ISSN 2227-4952 (Print), ISSN 2708-6143 (Online). Археологія і давня історія України, 2020, вип. 4 (37)
Lillie, M. C., Budd, C. E. Diet Isotope Analysis and Related Studies in Prehistoric Ukraine: Fact, Fiction and Fantasy
REFERENCES 
Ambrose, S. H. 1990. Preparation and characterization of 
bone and tooth collagen for isotopic analysis. Journal of Ar-
chaeological Science, 17, p. 431-451.
Ambrose, S. H., Norr, L. 1993. Experimental evidence for 
the relationship of the carbon isotope ratios of whole diet and 
dietary protein to those of bone collagen and carbonate. In: 
Lambert, J. B., Grupe, G. (eds.). Prehistoric Human Bone: Ar-
chaeology at the Molecular Level. New York: Springer, p. 1-37.
Ascough, P., Cook, G., Dugmore, A. 2005. Methodological 
approaches to determining the marine radiocarbon reservoir 
effect. Progess in Physical Geography, 29, 4, p. 532-547.
Balakin, S., Nuzhinyi, D. 1995. The origins of graveyards: 
the influence of landscape elements on social and ideological 
changes in prehistoric communities. Préhistoire Européenne, 
7, p. 191-202.
Brock, F., Higham, T., Ramsey, C. B. 2010. Pre-screening 
techniques for identification of samples suitable for radiocar­
bon dating of poorly preserved bones. Journal of Archaeologi-
cal Science, 37, p. 855-865.
Bronk Ramsey, C., Schulting, R. J., Goriunova, O. I., 
Bazaliiskii, V. I., Weber, A. W. 2014. Analyzing radiocarbon 
reservoir offsets through stable nitrogen isotopes and Baye­
sian modeling: a case study using paired human and faunal 
remains from the Cis-Baikal region, Siberia. Radiocarbon, 56, 
2, p. 1-11.
Buckley, M., Collins, M., Thomas-Oates, J., Wilson, J. C. 
2009. Species identification by analysis of bone collagen us­
ing matrix-assisted laser desorption/ionisation time-of-flight 
mass spectrometry. Rapid Communications in Mass Spec-
trometry, 23, p. 3843-3854.
Budd, C. 2015. Neolithic Anatolia and Central Europe: dis-
entangling environmental impacts from diet isotope studies. 
Doctoral dissertation. University of Oxford.
Budd, C., Lillie, M. C. 2020. The prehistoric popula­
tions of Ukraine: stable isotope studies of fisher-hunter-for­
ager and pastoralist-incipient farmer dietary pathways. In: 
Lillie, M. C., Potekhina, I. D. (eds.). Prehistoric Ukraine: 
From the First Hunters to the First Farmers. Oxford: Oxbow 
Books, p. 283-307.
Budd, C. E., Lillie, M. C., Potekhina, I. D. 2020 Continua­
tion of fishing subsistence in the Ukrainian Neolithic: diet iso­
tope studies at Yasinovatka, Dnieper Rapids. Archaeological 
and Anthropological Sciences, 12, 2, article id. 64.
Ciuk, K. (ed.). 2008. Mysteries of Ancient Ukraine: the re-
markable Trypilian Culture 5400—2700 BC. Canada: Royal 
Ontario Museum.
DeNiro, M. J. 1985. Post-mortem preservation and altera­
tion of in vivo bone collagen isotope ratios in relation to pale­
odietary reconstruction. Nature, 317, p. 806-809.
DeNiro, M. J., Epstein, S. 1978. Influence of diet on the 
distribution of carbon isotopes in animals. Geochimica Cos-
mochimica Acta, 42, p. 495-506.
Dolukhanov, P. 1979. Ecology and economy in Neolithic 
eastern Europe. London: Duckworth.
Dolukhanov, P. M. 2000. Alternative revolutions: hunter-
gatherers, farmers and stock-breeders in NorthWestern Pon­
tic area. In: Boyle, K., Renfrew, C., Levine. M. (eds.). Ancient 
Interactions: East and West in Eurasia. McDonald Institute 
Monographs. Cambridge, p. 13-24.
Dolukhanov, P. M., Khotinskiy, N. A. 1984. Human cul­
tures and the natural environments in the USSR during the 
Mesolithic and Neolithic. In: Vilechko, A. A. (ed.). Late Qua-
ternary Environments of the Soviet Union. Minneapolis: Uni­
versity of Minnesota, p. 319-327.
Gimbutas, M. 1956. The Prehistory of Eastern Europe. I: 
Mesolithic, Neolithic and Copper Age Cultures in Russia and 
the Baltic Area. ASPR Bulletin, 20. Cambridge.
Hedges, R. E., Reynard, L. M., 2007. Nitrogen isotopes and 
the trophic level of humans in archaeology. Journal of Archae-
ological Science, 34, 8, p. 1240-1251.
Henderson, R. C., 2015. Early life histories: a study of past 
childhood diet and health using stable isotopes and enamel 
hypoplasia. PhD Thesis. Oxford University.
Jacobs, K. 1993. Human Postcranial Variation in the 
Ukrainian Mesolithic-Neolithic. Current Anthropology, 34, 
p. 417-430.
Jacobs, K., 1994. Human Dento-Gnathic Metric Variation 
in Meolithic / Neolithic Ukraine: Possible Evidence of Demic 
Diffusion in the Dnieper Rapids Region. American Journal of 
Physical Anthropology, 95, p. 1-26.
Kadrow, S., Pokutta, D. A., 2016. The Verteba Cave: a 
subterranean sanctuary of the Cucuteni-Trypillia Culture in 
western Ukraine. Journal of Neolithic Archaeology, 18, p. 1-
21.
Karsten, J. K., Heins, S. E., Madden, G. D., Sokhat-
sky, M. P. 2014. The biological implications of the transition 
to agriculture in Ukraine: a study of enamel hypoplasias. In-
ternational Journal of Dental Anthropology, 27, 1—2, p. 16-
25.
Karsten, Dzh., Sokhatskiy, M. P., Kheyns, S. G., 
Medden, G. D. 2015a. Bioarkheologicheskiy analiz antropo­
logicheskogo materiala iz peshchery Verteba. Stratum plus, 
2, p. 121-144.
Karsten, J. K., Heins, S. E., Madden, G. D., Sokhat-
skyi, M. P. 2015b. Dental Health and the transition to agri­
culture in prehistoric Ukraine: a study of dental caries. Euro-
pean Journal of Archaeology, 18, 4, p. 562-579.
Kotova, N. 2018. Revisiting the Neolithic chronology of the 
Dnieper Steppe region with consideration of a Reservoir Ef­
fect for Human Skeletal Material. Sprawozdania Archeolog-
iczne, 70, p. 47-66.
Kozlowski, S. K. 1989. A Survey of Early Holocene Cul­
tures of the Western Part of the Russian Plain. In: Bonsall, C. 
(ed.). The Mesolithic in Europe. Edinburgh: J. Donald, p. 424-
441.
Krueger, H. W., Sullivan, C. H. 1984. Models for carbon iso­
tope fractionation between diet and bone. In: Turnland, J. R., 
Johnson, P. E. (eds.). Stable Isotopes in Nutrition. Washing­
ton DC: American Chemical Society Symposium Series, 258, 
p. 205-220.
Kruts, V. 2012. The Latest Stage of Development of the 
Tripolye Culture. In: Menotti, F., Korvin-Piotrovskiy, A. G. 
(eds.). The Tripolye Culture Giant Settlements in Ukraine: 
Formation, Development and Decline. Oxford: Oxbow Books, 
p. 230-253.
Ledogar, S. H., Karsten, J. K., Madden, G. D., Schmidt, R., 
Sokhatsky, M. P., Feranec, R. S. 2019. New AMS dates for 
Verteba Cave and stable isotope evidence of human diet in the 
Holocene forest-steppe Ukraine. Radiocarbon, 61, 1, p. 141-
158.
Lillie, M. C. 1996. Mesolithic and Neolithic populations of 
Ukraine: indications of diet from dental pathology. Current 
Anthropology, 37, 1, p. 135-142.
Lillie. M. C. 1997. Women and Children in Prehistory: 
Resource Sharing and Social Stratification at the Mesolithic-
Neolithic Transition in Ukraine. In: Moore, J., Scott, E. (eds.). 
Invisible People and Processes: Writing Gender and Childhood 
into European Archaeology. London: Leicester University, 
p. 213-228.
Lillie, M. C. 1998a. The Dnieper Rapids region of Ukraine: 
a consideration of chronology, dental pathology and diet at the 
Mesolithic—Neolithic transition. PhD Thesis. Sheffield Uni­
versity.
Lillie, M. C. 1998b. Cranial surgery dates back to Meso­
lithic. Nature, 391, p. 854.
Lillie, M. C. 1998c. The Mesolithic—Neolithic transition in 
Ukraine: new radiocarbon determinations for the cemeteries 
of the Dnieper Rapids region. Antiquity, 72, p. 184-188.
Lillie, M. C. 2003a. Tasting the Forbidden Fruit: gender 
based dietary differences among prehistoric hunter-gatherers 
of Eastern Europe? Before Farming, 2, 3, p. 1-16.
Lillie, M. C. 2003b. The Fruit and Nut Case: hunter-gath­
erer subsistence and egalitarianism in the riparian zone. In: 
Bevan, L., Moore, J. (eds.). Peopling the Mesolithic in a North-
ern Environment. British Archaeological Reports, Interna­
tional Series, 1157. Oxford: BAR Publishing, p. 59-68.
Lillie, M. C. 2020. Palaeopathology of the Prehistoric Pop­
ulations of Ukraine. In: Lillie, M. C., Potekhina, I. D. (eds.). 
Prehistoric Ukraine: From the First Hunters to the First Farm-
ers. Oxford: Oxbow Books, p. 235-281.
Lillie, M. C., Richards, M. P. 2000. Stable Isotope Analy­
sis and Dental Evidence of Diet at the Mesolithic-Neolithic 
Transition in Ukraine. Journal of Archaeological Science, 27, 
p. 965-972.
266 ISSN 2227-4952 (Print), ISSN 2708-6143 (Online). Археологія і давня історія України, 2020, вип. 4 (37)
Антропологія
Lillie, M. C., Jacobs, K. 2006. Stable isotope analysis of 
fourteen individuals from the Mesolithic cemetery of Vasily­
evka II, Dnieper Rapids region, Ukraine. Journal of Archaeo-
logical Science, 33, p. 880-886.
Lillie, M. C., Budd, C. 2011. The Mesolithic-Neolithic Tran­
sition in Eastern Europe: Integrating Stable Isotope Studies 
of Diet with Palaeopathology to Identify Subsistence Strate­
gies and Economy. In: Pinhasi, R., Stock, J. T. (eds.). Human 
Bioarchaeology of the Transition to Agriculture. Chichester: 
John Wiley & sons, p. 43-62.
Lillie, M. C., Richards, M. P., Jacobs, K. 2003. Stable Iso­
tope Analysis of Twenty-one Individuals from the Epipal­
aeolithic Cemetery of Vasilyevka III, Dnieper Rapids region, 
Ukraine. Journal of Archaeological Science, 30, p. 743-752.
Lillie, M. C, Budd, C. E., Potekhina, I. D. 2011. Stable iso­
tope analysis of prehistoric populations from the cemeteries 
of the Middle and Lower Dnieper Basin, Ukraine. Journal of 
Archaeological Science, 38, 1, p. 57-68.
Lillie, M. C., Budd, C. E., Potekhina, I. D., Hedges, R. E. M. 
2009. The Radiocarbon Reservoir Effect: New Evidence from 
the Cemeteries of the Middle and Lower Dnieper Basin, 
Ukraine. Journal of Archaeological Science, 36, 2, p. 256-264.
Lillie, M. C., Henderson, R., Budd, C. E., Potekhina, I. D. 
2016. Factors influencing the radiocarbon dating of human 
skeletal remains from the Dnieper river system: Archaeologi­
cal and stable isotope evidence of diet from the Epipalaeolithic 
to Eneolithic periods. Radiocarbon, 58, 4, p. 741-753.
Lillie, M. C., Budd, C. E., Potekhina, I. D. 2020. Radio­
carbon dating of sites on the Dnieper Region and Western 
Ukraine. In: Lillie, M. C., Potekhina, I. D. (eds.). Prehistoric 
Ukraine: From the First Hunters to the First Farmers. Oxford: 
Oxbow Books, p. 187-233.
Longin, R. 1971. New method of collagen extraction for ra­
diocarbon dating. Nature, 230 (5291), p. 241.
Madden, G. D., Kartsen, J. K., Ledogar, S. H., Scmidt, R., 
Sokhatsky, M. P. 2018. Violence at Verteba Cave, Ukraine: 
New insights into the Late Neolithic Intergroup Conflict. In-
ternational Journal of Osteoarchaeology, 28, p. 44-53.
Mallory, J. P. 1987. Editor’s Introduction. In: 
Telegin, D. Ya., Potekhina, I. D. Neolithic cemeteries and pop-
ulations in the Dnieper Basin. British Archaeological Reports, 
International Series, 383. Oxford: BAR Publishing, p. v-ix.
Mathieson, I., Alpaslan-Roodenberg, S., Posth, C., 
Szécsényi-Nagy, A., Rohland, N., Mallick, S., Olalde, I., 
Broomandkhoshbacht, N., Candilio, F., Cheronet, O. et al. 
2018. The Genomic History of Southeastern Europe. Nature, 
555, p. 197-203.
Minagawa, M., Wada, E. 1984. Stepwise enrichment of 15N 
along food chains: further evidence and the relation between 
δ15N and animal age. Geochimica et cosmochimica acta, 48, 5, 
p. 1135-1140.
Motuzaitë Matuzevičiűtë, G., Telizhenko, S. 2016. The 
first farmers of Ukraine: an archaeobotanical investigation 
and AMS dating of wheat grains from the Ratniv 2 site. Ar-
cheologia Lituana, 17, p. 100-111.
Müldner, G., Richards, M. P. 2005. Fast or feast: recon­
structing diet in later medieval England by stable isotope 
analysis. Journal of Archaeological Science, 32, 1, p. 39-48.
Nikitin, A. G. 2011. Bioarchaeological analysis of Bronze 
Age human remains from the Podillya Region of Ukraine. 
Interdisciplinaria Archaeologica: Natural Sciences in Archae-
ology, 2, 1/201, p. 9-14.
Nikitin, A. G., Sokhatsky, P., Kovaliukh, M. M., 
Videiko, M. Y. 2010. Comprehensive site chronology and an­
cient mitochondrial DNA analysis from Verteba Cave — a 
Trypillian culture site of Eneolithic Ukraine. Interdiscipli-
naria Archaeologica: Natural Science in Archaeology, 1, 1—2, 
p. 9-18.
Nikitin, A. G., Potekhina, I. D., Rohland, N., Mallick, S., 
Reich, D., Lillie, M. C. 2017. Mitochondrial DNA analysis of 
Eneolithic Trypillians from Ukraine reveals Neolithic farm­
ing genetic roots. PLoS ONE, 12, 2, e0172952.
O’Connell, T. C., Levine, M. A., Hedges, R. E. M. 2000. 
The importance of fish in the diet of central Eurasion peoples 
from the Mesolithic to the early Iron Age. In: Levine, M. A., 
Renfrew, A. C., Boyle, K. (eds.). Prehistoric Steppe Adaptation 
and the Horse. Cambridge: McDonald Institute for Archaeo­
logical Research, p. 253-68.
Philippsen, B. 2013. The freshwater reservoir effect in ra­
diocarbon dating. Heritage Science, 1, 1, p. 24.
Potekhina, I., Telegin, D. 1995. On the Dating of the 
Ukrainian Mesolithic-Neolithic Transition. Current Anthro-
pology, 36, 5, p. 823-826.
Potekhina, I., Lillie, M. C., Budd, C. E. 2014. Analiz pale­
odiyety za danymy stabilnykh izotopiv u populyatsiyakh epi­
paleolitu—eneolitu Serednioho ta Nyzhnioho Podniproviya. Is-
torychna antropolohiya ta bioarkheolohiya Ukrainy, I, s. 5-20.
Potekhina, I. D. 2018. Pokhodzhennya trypiltsiv iz pechery 
Verteba u svitli kraniolohiyi i arkheohenetyky. Mahisterium. 
Arkheolohichni studiyi, 70, с. 25-33.
Richards, M. P. 1998. Palaeodietary studies of European 
human populations using bone stable isotopes. D. Phil. Thesis. 
University of Oxford.
Richards, M. P. 2002. A brief review of the archaeological 
evidence for Palaeolithic and Neolithic subsistence. European 
Journal of Clinical Nutrition, 56, p. 1270-1278.
Sayle, K. L., Hamilton, W. D., Gestsdуttir, H., Cook, G. T. 
2016. Modelling Lake Mэvatn’s freshwater reservoir effect: 
Utilisation of the statistical program FRUITS to assist in 
the re-interpretation of radiocarbon dates from a cemetery at 
Hofstaрir, north-east Iceland. Quaternary Geochronology, 36, 
p. 1-11.
Schulting, R. J., Richards, M. P. 2001. Dating women and 
becoming farmers: new palaeodietary and AMS evidence from 
the Breton Mesolithic Cemeteries of Téviec and Hoлdic. Jour-
nal of Anthropological Archeology, 20, p. 314-344.
Schulting, R. J., Bronk Ramsey, C., Goriunova, O. I., 
Bazaliiskii, V. I., Weber, A.W. 2014. Analysing radiocarbon 
reservoir offsets with stable isotope data: a case study using 
paired human and faunal remains from the Cis-Baikal region, 
Siberia. Radiocarbon, 56, 3, p. 991-1008.
Schulting, R. J., Bronk Ramsey, C., Bazaliiskii, V. I., We­
ber A. W. 2015. Highly variable freshwater reservoir offsets 
found along the Upper Lena watershed, Cis-Baikal, southern 
Siberia. Radiocarbon, 57, 4, p. 1-13.
Schwarcz, H. P., Schoeninger, M. 1991. Stable isotope 
analysis in human nutritional ecology. Yearbook of Physical 
Anthropology, 34, p. 283-321.
Schoeninger, M., DeNiro, M. 1984. Nitrogen and carbon 
isotopic composition of bone collagen from marine and terres­
trial animals. Geochimica et Cosmochimica Acta, 48, p. 625-
639.
Sealy, J. C., Johnson, M., Richards, M., Nehlich, O. 2014. 
Comparison of two methods of extracting bone collagen for 
stable carbon and nitrogen isotope analysis: comparing whole 
bone demineralization with gelatinization and ultrafiltration. 
Journal of Archaeological Science, 47, p. 64-69.
Telegin, D. Ya. 1986. Derievka: a settlement and cemetery 
of Copper Age horse keepers on the middle Dnieper. British 
Archaeological Report, S287. Oxford: BAR Publishing.
Telegin, D. Ya., 1987. Neolithic cultures of the Ukraine 
and adjacent areas and their chronology. Journal of World 
Prehistory, 1 (3), p. 307-331.
Telegin, D. Ya., Potekhina, I. D. 1987. Neolithic cemeteries 
and populations in the Dnieper Basin. British Archaeological 
Report, S383. Oxford: BAR Publishing.
Telegin, D. Ya., Titova, E. N. 1993. La Zone des Steppes. 
In: Kozlowski, J (ed.). Atlas du Néolithique européen. 1: 
L’Europe orientale. (Études et Recherches Archéologiques de 
l’Université de Liège, 45. Liège, p. 463-494.
Telegin, D. Ya., Zaliznyak, L. L. 1975. Raskopki na 
Igren’skom poluostrove. Arkheologicheskiye otkrytiya 1974 g., 
s. 358-359.
Telegin, D. Ya., Potekhina, I. D., Kovaliukh, M. M., Lil-
lie, M. C. 2000. Chronology of the Mariupol-type cemeteries 
and the problem of the periodisation of the Neolithic to Cop­
per Age cultures of Ukraine. Radiocarbon & Archaeology, 1, 
1, p. 59-74.
Telegin, D. Ya., Potekhina, I. D., Lillie, M. C., Kova-
liukh, M. M. 2002. The chronology of the mariupol-type cem­
eteries of Ukraine re-visited. Antiquity, 76, p. 356-363.
Telegin, D. Ya., Lillie, M. C., Potekhina, I. D., Kova-
liukh, M. M. 2003. Settlement and economy in Neolithic 
Ukraine: a new chronology. Antiquity, 77 (279), p. 456-470.
Tieszen, L. L., Fagre, T. 1993. Effect of diet quality 
and composition on the isotopic composition of respira­
267ISSN 2227-4952 (Print), ISSN 2708-6143 (Online). Археологія і давня історія України, 2020, вип. 4 (37)
Lillie, M. C., Budd, C. E. Diet Isotope Analysis and Related Studies in Prehistoric Ukraine: Fact, Fiction and Fantasy
tory CO2, bone collagen, bioapatite, and soft tissues, In: 
Lambert, J. B., Grupe, G. (eds.). Prehistoric Human Bone: Ar-
chaeology at the Molecular Level. New York: Springer, p. 121- 
155.
Van Klinken, G. J. 1999. Bone collagen quality indicators 
for palaeodietary and radiocarbon measurements. Journal of 
Archaeological Science, 26, 6, p. 687-695.
M. C. Lillie, C. E. Budd 
DIET ISOTOPE ANALYSIS  
AND RELATED STUDIES  
IN PREHISTORIC UKRAINE:  
FACT, FICTION AND FANTASY 
The authors consider scientific studies of Ukrainian 
skeletal material across the Epipalaeolithic to Eneo­
lithic periods and offer some observations in relation 
to the efficacy of studies undertaken by different re­
searchers. Firstly, the authors summarize the results 
of their own research since the original research under­
taken by Lillie in the early 1990s, and present period 
based overviews (fig. 1—3) which discuss the nature 
of the evidence, the fact that fish remains important 
across the periods studied. The data also highlights 
the fact that by the Eneolithic period different culture 
groups are following distinct subsistence strategies. 
This is obviously marked by western dietary pathways 
linked to the integration of agro-pastoralism (and asso­
ciated to presumed Trypillia farming groups at Verte­
ba Cave), and those of the eastern hunter-fisher-forag­
ers in the Dnieper region at Igren VIII and Molukhov 
Bugor.
Interestingly the chronological separation between 
these two sites is also linked to dietary variability. 
At the earlier site of Igren VIII there is diet isotope 
evidence for a relatively heavy reliance on freshwa­
ter resources as ca. 4300—4000 cal BC, whilst at the 
latter site of Molukhov Bugor, at 3950—3700 cal BC, 
a reduction in the reliance on freshwater resources 
is in evidence. This is accompanied by evidence for a 
broader spectrum approach to the exploitation of the 
wild resources in and around the Dnieper Rapids re­
gion. Radiocarbon dating is shown to be affected by the 
FRE at the sites in and around the Dnieper system. 
This is clearly not the case at Verteba Cave because 
the freshwater reservoir effect is not associated with 
dietary pathways that place a reliance on terrestrial 
resources.
The authors discuss the dating (fig. 4—6) and mobil­
ity and dietary isotope studies that they undertook at 
this location and contextualize these by comparison to 
the work of other researchers. It is suggested that some 
issues occur in relation to the different research groups 
activities at Verteba Cave, and the fact that there is a 
clear need for a more considered approach to the data 
presented by these other groups is highlighted. It could 
be argued that a lack of detailed knowledge and col­
laboration occurs despite the fact that there are clear 
overlaps between research agendas. The authors con­
clude with a call for targeted multi-disciplinary analy­
ses aimed at whole cemetery studies in order to further 
enhance our understanding of socio-economic and so­
cietal developments during the early to mid-Holocene 
in Ukraine.
Keywords: Ukraine, prehistory, dietary isotope 
analysis, δ13C, δ15N, AMS dating, freshwater reservoir 
effects.
М. К. Ліллі, Ч. Е. Бад 
іЗотопниЙ діЄтниЙ АнАліЗ тА 
поВ’ЯЗАні З ниМ дослідженнЯ 
перВісної історії УкрАїні: ФАк-
ти, ВигАдки тА ФАнтАстикА 
Автори розглядають результати вивчення скелет­
ного матеріалу періоду від епіпалеоліту та енеоліту та 
наводять деякі спостереження щодо ефективності до­
сліджень, проведених різними вченими. Перш за все, 
автори узагальнюють результати досліджень, які про­
водились М. Ліллі з колегами з початку 1990-х рр., та 
проводять огляд нових результатів (рис. 1—3), які під­
тверджують той факт, що риба залишалася важливим 
продуктом харчування протягом названих періодів. 
Отримані дані також підкреслюють той факт, що в 
епоху енеоліту різні культурні групи дотримувалися 
різних харчових стратегій. Це, очевидно, визначаєть­
ся західними впливами та інтеграцією землеробства 
і скотарства (трипільська фермерська група з печери 
вертеба), та традиціями мисливців-рибалок-збирачів 
з Ігрені VIII та Молюхового бугра у Подніпров’ї.
Цікаво, що хронологічна відмінність між цими дво­
ма пам’ятками супроводжується і харчовою мінливіс­
тю. На старшій пам’ятці Ігрень VIII отримані ізотопні 
дані про відносно високу залежність від прісноводних 
ресурсів у період 4300—4000 cal BC, тоді як на піз­
нішій пам’ятці Молюхів бугор, дані свідчать про змен­
шення залежності від прісноводних ресурсів протягом 
3950—3700 cal BC. Ці результати супроводжуються 
доказами використання більш широкого спектру 
природних ресурсів у регіоні Дніпровських порогів та 
навколо нього. Показано також, що на радіовуглецеві 
дати на пам’ятках у басейні Дніпра впливає прісно­
водний резервуарний ефект. Цього не спостерігається 
на матеріалах з печери вертеба, оскільки ефект пріс­
новодного резервуару не пов’язаний з дієтичними лан­
цюгами, які залежать від наземних ресурсів.
Отримані авторами результати датування (рис. 4—
6), дослідження мобільності та дієтних ізотопів обго­
ворюються у контексті результатів інших дослідників. 
висловлюється припущення, що діяльність різних 
дослідницьких груп у печері вертеба породжує низку 
проблем, що викликає необхідність продуманішого під­
ходу до даних, представлених іншими групами. Можна 
стверджувати, що незважаючи на спільну тематику і 
близькі програми, окремим дослідницьким групам бра­
кує детальних знань та співпраці. Стаття завершується 
закликом до цілеспрямованого мультидисциплінарно­
го аналізу при дослідженні всіх могильників з метою 
подальшого вдосконалення нашого розуміння соціаль­
но-економічних та суспільних подій протягом раннього 
та середнього голоцену на території України.
ключові слова: Україна, передісторія, аналіз 
дієтних ізотопів, δ13C, δ15N, датування AMS, прісно­
водний резервуарний ефект.
Одержано 10.07.2020
БАд Челсі, Доктор філософії, Постдокторант уні­
верситету м. Умеа, SE 901-87, Швеція.
bUDD Chelsea, PhD, Postodcotral Research Fellow, 
Umeå University, SE 901-87, Sweden.
ORCID: 0000-0001-8527-9669, e-mail: Chelsea.budd@umu.se.
ліллі Малкольм, Доктор філософії, Професор уні­
верситету м. Умеа, SE 901-87, Швеція.
LILLIE Malcolm, PhD, Professor, Umeå University, 
SE 901-87, Sweden.
ORCID: 0000-0002-6054-3651, e-mail: malcolm.lillie@
umu.se.