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Journal of Archaeological Science (1999) 26, 717722

Article No. jasc.1998.0387, available online at http://www.idealibrary.com on

Stable Isotope Evidence for Similarities in the Types of Marine


Foods Used by Late Mesolithic Humans at Sites Along the
Atlantic Coast of Europe
M. P. Richards and R. E. M. Hedges
Research Laboratory for Archaeology and the History of Art, 6 Keble Road, Oxford OX1 3QJ, U.K.

Human bone collagen stable isotope analysis of humans from coastal Mesolithic sites in Scotland, Denmark, France
and Portugal indicates the importance of marine foods in the diet. We define the expected human 13C and 15N values
of 100% marine and 100% terrestrial diets and conclude that at most sites isotope variability is due to differing
proportions of these defined marine and terrestrial diets, rather than due to differences in the actual types of marine and
terrestrial foods exploited. By comparing the European human values with marine faunal values, and values of
marine-diet humans from North America, we propose that the marine component of human diet in the Late European
Mesolithic was based mainly on marine fish, with only minor contributions from shellfish or marine mammals.
 1999 Academic Press

Keywords: STABLE ISOTOPES, CARBON, NITROGEN, MESOLITHIC, PALAEODIET, TE


u VIEC, HO
} EDIC,
PORT AU CHOIX, MARINE RESOURCES.

Introduction humans, that eat plants or that eat other animals, has
a 13C value of around 20 (1) . The bone

T
here are many Late Mesolithic sites along the collagen of animals at a higher trophic level tends to be
Atlantic coast of Europe that contain evidence, slightly heavier in 13C, and there is some additional
in the form of midden deposits of marine shells, variation due to climate and other effects. The 15N
and fish and marine mammal bones, for the human value depends primarily on the trophic level of the
consumption of marine foods. We discuss here how animals diet, but also on the general interaction
stable isotope analysis of surviving collagen in human between soil, vegetation and climate; 15N ranges
bone from these sites can complement faunal analysis, from 4 to 10 in prehistoric European humans
can help to determine the relative importance, in (Schoeninger, DeNiro & Tauber, 1983; Murray &
nutritional terms, of marine and terrestrial resources Schoeninger, 1988; Richards & Hedges, unpubl.
and may indicate what some of these resources might data).
be. We present new results from two Late Mesolithic Marine food chains are more complicated and less
sites (in Brittany), and we include in the discussion well defined isotopically. Photosynthesis of dissolved
published results from Scotland, Portugal and CO2 by phytoplankton results in less fractionation
Denmark. These results are compared with published than in C3 plants, and carbon in the food web is
and unpublished measurements from human bone generally isotonically heavier by about 8. However,
collagen from populations on the east and west coasts global variations can be quite large, and are correlated
of North America, and are also discussed in relation to with sea temperature and therefore dissolved CO2
stable isotope analyses of marine fauna. content. Isotope values of humans eating a 100%
marine food diet are not so easily determined as for a
100% terrestrial food diet, but it is possible to use
Isotope Values in Foodwebs marine mammals as a comparison. In particular, sea
This vast topic has been reviewed in part numerous mammals from the North Atlantic have similar colla-
times (e.g., Ambrose, 1993; DeNiro, 1987; Owens, gen 13C values (of 12) to those from the west
1987; Schwarcz & Schoeninger, 1991; van der Merwe, coast of North and South America, and this similarity
1982) and we summarize the relevant points here. The of collagen 13C marine signature is also seen in other
assumptions and approximations involved in the examples (Table 1). (Note that comparison must be
analysis we make are identified separately. between collagen values, since flesh 13C values are
In the regions under consideration in this paper, offset from collagen values.) The enrichment of 15N at
terrestrial food chains are based on photosynthesis by higher trophic levels is particularly apparent and well
C3 plants. The bone collagen of animals, including known for marine systems, partly because the marine
717
03054403/99/060717+06 $30.00/0  1999 Academic Press
718 M. P. Richards and R. E. M. Hedges

Table 1. Average 13C and 15N values of modern and archaeological marine fauna from various locations around the world

Class of fauna Material 13C1 15N1 N Location of samples, and source of data

Sea lions Bone coll. 12708 18015 22 4 M CA(1), 15 A CA(2), 3 A PE(2)


Seals Bone coll. 12313 17021 13 (10) 2 M CA(1), 1 M UN (1), 1 M NWE (1),
2 BC M+A(3), 3 M SA(4), 3 A NWE(2)
Whales (A) Bone coll. 12911 16508 5 1M AK(1), 4 M CA(1)
Dolphins/porpoises Bone coll. 12810 15709 12 12 M CA(1)
Fish (C) Bone coll. 12816 12821 3 3 M SC(1)
Fish (C) Flesh 17205 14528 9 (7) 7 M GB(5), 2 M SA(4)
Whales (B) Bone coll 14514 13713 12 (10) 3 M AK(1), 7 M CA(1), 2 M SA(4)
Fish (B) Bone coll 13212 13409 21 7 M SC(1), 14 M EQ(6)
Fish (B) Flesh 16719 12619 45 4 M UN(7), 5 M UN(8), 36 M GB(5)
Cephalopods Flesh 16704 12623 5 3 M EQ(6), 2 M GB(5)
Walruses Bone coll 13903 12302 3 2 M AK(1), 1 M NWE(1)
Fish (A) Flesh 19319 11519 11 5 M BC(9), 6 M GB(5)
Crustaceans Flesh 1572 10723 14 (13) 10 M GB(5), 3 M EQ(6), 1 M SA(4)
Shellfish Flesh 15920 8813 45 (36) 13 M EQ(6), 3 M UN (8), 20 M GB(5), 9 M SA(4)

*Whales (A) consume primarily molluscs, arthropods and plankton, while Whales (B) consume mainly fish. Fish (A) are planktivores, Fish (B)
are opportunistic generalists, or have an unknown diet, and Fish (C) are piscivores.
Bone coll.=bone collagen.
Numbers in parentheses refer to N values for 13C averages, where this differs from the 15N.
M=Modern, A=Archaeological, CA=California, PE=Peru, UN=Unknown, BC=British Columbia, Canada, NWE=Northwest Europe,
SA=South Africa, AK=Alaska, GB=Georges Banks, NW Atlantic, EQ=Equador. Reference numbers: (1) Schoeninger & DeNiro (1984), (2)
DeNiro (1985), (3) Richards (unpubl. data), (4) Sealy et al. (1987) only 15N values, (5) Fry (1988), (6) van der Merwe, Lee-Thorp & Raymond
(1993) (7) Bonsall et al. (1997), (8) Minagawa (1992), (9) Hobson (1990).

20.0
19.0
18.0
17.0
16.0 Sea Lions
Whales (B) N = 22
15.0
N = 5 Seals
14.0 Dolphins/ N = 13
15N ()

Porpoises
13.0 N = 12
12.0 Whales (A)
Walrus
N=3 N = 12
11.0 Fish (C)
Fish (B)
10.0 N = 66 N = 12
Cephalopods
9.0 Fish (A) N=5
N = 11
8.0 Crustaceans
N = 14
7.0 Shellfish
6.0 N = 45
5.0
Species
15
Figure 1. Average N values of modern and archaeological marine fauna. The sources of the data are given in Table 1. Fish (A) are
planktivores, Fish (B) are opportunistic generalists, or have unknown diets, and Fish (C) are piscivores. Whales (A) eat plankton, molluscs and
arthropods, while Whales (B) are primarily piscivores.

food chain has many more trophic levels than the 15N (1) the range involved, (2) how animals at differ-
terrestrial one (Schoeninger & DeNiro 1984). Figure 1 ent trophic levels differ, and (3) that there is no
& Table 1 list faunal 15N data (note that collagen and appreciable geographic difference in 15N values for
flesh 15N are approximately the same), and show in animals at similar trophic levels. If humans had a diet
Marine Foods Used by Late Mesolithic Humans in Europe 719

Marine HIGH 13C values close to 20 indicate a diet with almost
24
trophic level
carnivore no (<5%) marine protein, and 13C values close to
22 12 indicate a diet of almost 100% marine protein.
20 The measurement of 15N can be used to further define
18
the types of marine protein in the diets, as the 15N
15N ()

16
14 helps to define the trophic level of the marine foods
12 Terrestrial
carnivore
consumed.
10
8 Marine LOW
trophic level
6
4
carnivore
Assumptions and Approximations Involved in
2 Terrestrial
herbivore
the Analysis
0
22 21 20 19 18 17 16 15 14 13 12 11 10 It may be helpful to list the assumptions and approxi-
13
C () mations used here.
Figure 2. Four (theoretical) extreme human dietary types, and their (1) That human bone collagen composition refers to
associated bone collagen stable isotope values. diet averaged over the 510 years prior to death.
(2) That the collagen 13C value is determined by the
weighted average of the dietary protein 13C, plus
of only marine food, they would be expected to have an offset, and is not affected by other aspects of
bone collagen values enriched by approximately 34 the diet. (The offset is 35 between collagen and
in 15N above that of the diet. flesh, but 01 between human collagen and the
It is therefore possible to identify four extreme collagen value of the animal consumed. However,
values for human bone collagen stable isotopes, which this offset does not enter directly into the
correspond to four extreme types of diet (Figure 2). quantitative considerations here.)
(3) That the collagen 15N value is a weighted aver-
age also of the dietary protein 15N, with a offset
A Framework for the Analysis of Marine and of +3, and this offset is not altered by the
Terrestrial Diets nutritional quality of the diet.
(4) That human collagen from a diet based on a
Human bone collagen should have an isotopic value terrestrial C3 food web has a 13C value that may
within the field defined by the extreme points given in be approximated as 20 for the regions
Figure 2, but to what extent does this actually define studied. (This removes the complexities of con-
possible human diets? sidering diet-collagen offsets. It is probably not a
Consider carbon first. In effect there are only two very accurate approximation, but we believe is
sources, marine and terrestrial. We assume that inter- sufficient for our purposes here.)
mediate sources, for example, sea birds, otters, eels, (5) That human collagen from a wholly marine diet
etc., which may feed on either marine or terrestrial has a 13C value that may be approximated as
food chains, are not significant dietary components 121.
and can be neglected. If a human eats a combination of (6) That human collagen 15N ranges from 4 to 10
marine and terrestrial components, the resulting bone for terrestrial diets and from 10 to 22 for
collagen isotope value may not be proportional to the marine diets, depending on the trophic level of the
relative carbon in each component. In fact there is food protein.
considerable evidence from controlled feeding exper-
iments on mice and rats that with an adequate protein
diet, bone collagen values are proportional to the
carbon content of the protein components in the diet
The Data Set
(Ambrose & Norr, 1993; Tieszen & Fagre, 1993). This Figure 3 shows hitherto unpublished and recently
is equivalent to the notion that collagen carbon is published collagen values for human bone collagen
derived only from ingested protein carbon, and that all from three Late Mesolithic sites; Oronsay, a small
protein is treated equally, which is not necessarily true. island in the Inner Hebrides, Scotland (Richards &
A similar assumption applies to nitrogen, that all Mellars 1998), and Teviec and Hoedic, two small
collagen nitrogen comes equally from ingested protein, islands off the south coast of Brittany, France
but the validity of this assumption is more secure, (Richards & Schulting, unpubl. data). Details of
because protein is the only significant dietary source of sample preparation for samples from these last two
nitrogen, and amino acids do not differ greatly in 15N sites are given in Appendix 1. Also plotted on this
amongst themselves (Hare et al., 1991). figure are data from humans from several Portuguese
It follows from this that the proportion of marine sites (Lubell et al., 1994), and average 13C and
protein in the dietary protein can be easily estimated by 15N values of Mesolithic humans from Denmark
linear interpolation using the bone collagen 13C value. (Schoeninger et al., 1983).
720 M. P. Richards and R. E. M. Hedges

22.0 Table 2. Average bone collagen 15N values of humans whose dietary
protein was almost exclusively from marine sources, as the 13C values
20.0 _14
Seal
18.0
16.0 Otter Geographic region 15N1 N
15N ()

14.0
12.0 Saithe Mesolithic Europe 14513 15
10.0 Crustaceans Newfoundland 20306 19
8.0 British Columbia 18613 29
Shellfish
California 17018 8
6.0
4.0
20.0 19.0 18.0 17.0 16.0 15.0 14.0 13.0 12.0 11.0 The Mesolithic European data consist of five values from Oronsay
13 (Richards & Mellars, 1998), three values from Hoedic (Richards &
C () Schulting, unpubl. data), and seven values from Denmark, taken
Figure 3. Stable isotope values of human bone collagen from five from Figure 1(A) in Schoeninger et al. (1983). The Port au Choix
Late Mesolithic sites along the Atlantic coast of Europe. The data are a combination of measurements made at Oxford by M. R.
Oronsay values are from Richards & Mellars (1998), the Portuguese on samples provided by J. Jelsma (Richards & Jelsma, unpubl: data)
values are from Lubell et al. (1994), the Teviec and Hoedic values are and measurements made at Groningen (Jelsma, unpubl: data). Data
from Richards & Schulting (unpubl. data), and the Danish values are from British Columbia are taken from Figure 5 in Schwarcz (1991).
taken from Schoeninger et al. (1983). The names of various marine The Californian data are taken from Figure 4 in Walker & DeNiro
organisms are also plotted at a level corresponding roughly to their (1986).
average 15N value. , Teviec; , Hoedic; , Portugal Mesolithic;
, Oronsay; , Denmark.
15N values of humans from various (unnamed) pre-
historic coastal sites in British Columbia to be indica-
22.0 tive of a primarily salmon diet. Walker & DeNiro
20.0 (1986) measured the isotope values of humans from
prehistoric and historic coastal, island and inland
18.0
sites in the Santa Barbara channel area in southern
16.0 California. Ethnographic accounts of early European
N ()

14.0 explorers in this region notes that the indigenous


peoples obtained nearly all of their food through
15

12.0
fishing (Walker & DeNiro, 1986: 57). In particular,
10.0 there are large schools of small fish in this region that
8.0 support larger fish and sea mammals. They conclude
6.0 that the isotope values, particularly the 15N values,
18.0 17.0 16.0 15.0 14.0 13.0 12.0 11.0 suggest a diet similar to the local marine mammals
13C () (whose isotopes they also measured) and heavily
dependent on fish, rather than shellfish or marine
Figure 4. Bone collagen 13C and 15N values of humans from
various archaeological sites in North America. The Newfoundland mammals. C4 plants were not a part of the diet of
data are from Richards & Jelsma (unpubl. data) and Jelsma (in humans or fauna consumed by humans at these
press), the British Columbian data are taken from Figure 5 in Californian sites (Walker & DeNiro, 1986), or indeed
Schwarcz (1991), and the Californian data are taken from Figure 4 at the Newfoundland, British Columbia and European
in Walker & DeNiro (1986). , Newfoundland; +, British Columbia;
, California.
sites.

Figure 4 is a plot of new and previously published


data from North America for a comparison. These
Discussion
data are from humans of a much greater range of date, In Table 2 the average bone collagen 15N values of
but for which there is strong circumstantial evidence humans with predominantly marine diets (13C
for their diet. The humans from the Port au Choix site d14) from the sites discussed above are given.
in Newfoundland, Canada (c. 32004300 ; Orban, The 15N for humans with a predominantly marine
1989) were primarily seal hunters. This assertion is diet at the Port au Choix site in Newfoundland is
based on ethnographic accounts of more recent peoples particularly high (20306), and is consistent with
living in the same region, as well as on the large the expected marine diet based on seals. The British
numbers of seal bones and artefacts associated with Columbian values, 15N=18513, are higher than
seal hunting found at the site (Jelsma, in press). one would expect for a marine diet based mainly on
Humans from coastal British Columbia are known salmon, for which one might expect human values of
from ethnographic accounts to be primarily salmon about 15N=17. Instead, the values suggest the
fishers, and Schwarcz (1991) and Chisholm, Nelson & inclusion of some marine mammals, like seals, in the
Schwarcz (1983) assumed this pattern of subsistence to diet. However, there is a large spread in these values, so
be similar in the past, and interpreted the 13C and it is probable that there is not a single type of marine
Marine Foods Used by Late Mesolithic Humans in Europe 721

food diet at coastal sites in British Columbia. As the correlate well enough to produce a straight line. The
data are from a variety of sites and time periods that European Mesolithic 13C and 15N data, with the
are not specified, this is probably the case. The 15N exception of the two circled points in Figure 3, also
values of the marine-diet humans from southern correlate well. These correlations at these different sites
California average 1718, and suggest a marine are difficult to explain in any other simple way than as
food diet consisting of fairly high trophic level fauna, a two-component mixture of a consistent marine and a
such as piscivorous fish, which is consistent with consistent terrestrial diet, the proportions of which
Walker & DeNiros conclusions given above. might vary. If each type of diet contains only one
When the 13C and 15N values of humans from component (in terms of protein 15N) then any dietary
both the Californian and the Newfoundland sites are variation at all would have to produce a straight line,
plotted each data set lies along a well-correlated line and such a line is an argument for there being only one
(these lines are drawn in Figure 4). This indicates that source of protein exploited (or rather, for protein at
the differences in 13C and 15N values between indi- only one trophic level being exploited). However, it
viduals in the same populations is due to diets of could be that a mixture of resources is exploited, but
differing proportions of marine and terrestrial foods that this mixture does not change from one individual
within those populations, but not differences in the to another.
types of those marine and terrestrial foods consumed.
This aspect will be returned to after considering the Summary
European data.
By comparison, the European data are more scat- Humans buried at European Mesolithic coastal sites
tered, show a wider range of 13C and 15N values, and generally obtained more than 50% of their protein
have generally lower 15N values than the humans from marine resources, and in some cases up to 90%.
from North America (Figure 3). The highest values are They appear to have eaten marine food characterized
found at Oronsay, at 1516 they are some 67 by a lower trophic level than comparative archaeologi-
lower than those for the supposed seal-eaters of cal populations in both coasts of North America.
Port au Choix, and are consistent with the marine However, this does not mean they mainly relied on
dietary protein being mainly fish of a generalist shellfish, and indeed for the most part shellfish prob-
diet, or alternatively a mixture of seals and shellfish. ably constituted much less than 50% of their marine
The few humans from Hoedic with marine diets protein. Finally, the variation between individuals fre-
(13C d14) have similar 15N values, of about quently shows an interesting pattern that implies they
16, and therefore also indicate dietary marine differed mainly in the relative proportions of terrestrial
protein with an average value of about 13. and marine foods, while the composition of those
The humans from Teviec had more mixed marine foods was rather conservatively maintained. The vari-
and terrestrial protein diets, with more negative 13C ous explanations advanced for differences in the
values. However, it is possible to extrapolate the Teviec marine to terrestrial food proportions are worth pur-
values to estimate what the 15N values would be with suing since they deal with important cultural aspects of
more marine protein in the diet (if the 13C values were resource exploitation, such as seasonality.
d14), and the resulting 15N values would be
similar to those from Hoedic. If a similar estimation
was made from the Portuguese data the resulting 15N References
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