Isotope and faunal evidence for high levels of freshwater fish consumption by Late Glacial humans at the Late Upper Palaeolithic site of Šandalja II, Istria, Croatia

Journal of Archaeological Science 61 (2015) 204e212

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Isotope and faunal evidence for high levels of freshwater fish consumption by Late Glacial humans at the Late Upper Palaeolithic
site of Sandalja II, Istria, Croatia M.P. Richards a, b, *, I. Karavani c c, P. Pettitt d, P. Miracle e a

Department of Anthropology, University of British Columbia, 6303 NW Marine Drive, Vancouver, BC, V6T1Z1, Canada Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, Leipzig, D-04103, Germany Department of Archaeology, Faculty of Humanities and Social Sciences, University of Zagreb, Ivana Lu
cica 3, 10000, Zagreb, Croatia d Department of Archaeology, Durham University, Dawson Building, South Road, Durham, DH1 3LE, UK e Department of Archaeology, University of Cambridge, Downing Street, Cambridge, CB2 3DZ, UK b c

a r t i c l e i n f o

a b s t r a c t

Article history: Received 17 September 2014 Received in revised form 9 June 2015 Accepted 10 June 2015 Available online 19 June 2015

Here we report on isotope and faunal evidence for intensive use of freshwater resources by Late Upper
Palaeolithic humans from the Sandalja II site in Croatia. Carbon and nitrogen bone collagen isotopic analysis of humans and fauna from the site indicate that the main protein source in human diets at this time was freshwater fish, which is in contrast to the vertebrate remains that show a high abundance of large terrestrial herbivores from the Late Upper Palaeolithic levels at the site. These data add to the growing body of research that shows an increasing intensification in the use of aquatic resources in Europe towards the end of the Pleistocene. © 2015 Elsevier Ltd. All rights reserved.

Keywords: Late Upper Palaeolithic Palaeodiet Isotopes Freshwater fish

1. Introduction Isotope analysis of past diets in Palaeolithic Europe has largely focussed on contrasting the dietary adaptations of Middle Palaeolithic Neanderthals and early Upper Palaeolithic modern humans (Richards and Trinkaus, 2009) and studies of Late Upper Palaeolithic material (ca. 15,000e12,000 cal BP) have been few and far between. Despite this, the Late Upper Palaeolithic is a potentially important period to capture isotopically, as it may well be the period in which one can see the earliest evidence for increasing dietary specialisation and resource intensification, e especially of aquatic resources e which became more frequent in the subsequent Holocene, especially in the coastal Western European Mesolithic, where humans had diets of close to 100% marine foods in Denmark and the UK (e.g. Richards et al., 2003a,b; Fischer et al., 2007).

* Corresponding author. Department of Anthropology, University of British Columbia, 6303 NW Marine Drive, Vancouver, BC, V6T 1Z1, Canada. Tel.: þ1 604 822 3503. E-mail address: [email protected] (M.P. Richards). http://dx.doi.org/10.1016/j.jas.2015.06.008 0305-4403/© 2015 Elsevier Ltd. All rights reserved.

A handful of isotope studies on Late Upper Palaeolithic (ca. 15,000e12,000 BP) human remains from Western Europe have been undertaken, deriving from the UK (Richards et al., 2000, 2005), France (Drucker et al., 2005; Naito et al., 2013), Spain  et al., 2009) and Italy (Craig et al., 2010; Francalacci, (Garcia-Guixe 1988; Mannino et al., 2011a,b; 2012, Gazzoni et al., 2013). To date there have been no applications to humans from this time period across much of the continent and particularly in Eastern or South East Europe. In this study we sampled humans and fauna from the Late Upper
Palaeolithic levels of Sandalja II, Istria, Croatia. Of interest here was the potential use of plant food resources in this temperate environment, as well as marine and freshwater foods. Previous zooarchaeological analysis of the upper Late Palaeolithic levels at this site revealed the presence of freshwater fish remains (Miracle, 1995; Paunovi c, 1984, 1987), although the vertebrate assemblage is overwhelmingly dominated by large mammals (Miracle, 1995, 1996, 2007). Our study was therefore undertaken to see if the isotope evidence supports an argument for the intensive use of freshwater fish by humans during the Late Upper Palaeolithic at this site, or whether it was relatively uncommon. Either way we intended to add to the increasing body of literature on human

M.P. Richards et al. / Journal of Archaeological Science 61 (2015) 204e212

dietary adaptations during the final stage of the Pleistocene in Europe. 2. Previous isotope studies Carbon and nitrogen isotopic values of bone collagen are directly related to the carbon and nitrogen isotope values of dietary protein consumed (DeNiro and Epstein, 1978, 1981, Schoeninger et al., 1983, 1984). Adult human bone collagen is an average of many years of dietary protein intake, and thus reflects the overall major dietary protein sources over perhaps the last 10e20 years of life (Hedges et al., 2007). There are many reviews of isotope analysis for dietary reconstruction (e.g. Lee-Thorp, 2008) and it has become a well-established technique, so we will not review the background here. In this study we are using carbon (d13C) and nitrogen (d15N) bone collagen isotope analysis to look for the relative contribution of marine vs. terrestrial protein in long-term human diets, as well as the relative amounts of plant vs. animal protein and, if detectable, the consumption of freshwater resources. Previous isotope studies of Late, or Final Upper Palaeolithic (ca. 15,000e12,000 cal BP) humans from the UK revealed a range of dietary adaptations ranging from dependence on the hunting of terrestrial ungulates for three individuals at Gough's and Sun Hole Caves in Somerset (Richards et al., 2000; Jacobi and Higham 2009), to the supplementing of terrestrial foods with marine foods, consisting of perhaps 25% of dietary protein, for three individuals from the site of Kendrick's Cave in north Wales (Richards et al., 2005) (Table 1). More applications of isotope analysis to Late Upper Palaeolithic humans have been undertaken in Italy (Table 1). These comprise human remains from the Sicilian sites of Grotta d'Oriente (Mannino et al., 2012), Grotta Addaura Caprara and Grotta di San Teodoro (Mannino et al., 2011a, b), and the mainland sites of Grotta del Romito (Craig et al., 2010), Ripero Tagliente (Gazzoni et al., 2013) and Arene Candide (Francalacci, 1988). Most of these Italian Late Upper Palaeolithic humans had d13C values centred around 20‰

205

and d15N values close to 9‰, indicating that the source of their dietary protein was largely terrestrial, with little input from aquatic resources, despite the close proximity of a number of the sites to the coast. Three individuals did have isotopic evidence for regular aquatic food consumption. A single individual from the site of Grotta del Romito (Romito 9) had isotope values indicative of marine food consumption, probably in addition to regular freshwater fish consumption, with a d13C value of 18.9‰ and a relatively high d15N value of 12.4‰ (Craig et al., 2010). Similarly, a single individual from the site of Ripero Tagliente had a d13C value of 18.4‰, and an elevated d15N value of 13.0‰, also indicating likely marine food consumption in addition to freshwater fish consumption (Gazzoni et al., 2013). One individual from San Teodoro 1 had a terrestrial d13C value of 20‰, but an elevated d15N value of 12.5‰. In this case the authors concluded that the San Teodoro 1 individual likely consumed a significant amount of freshwater fish in addition to terrestrial resources, but the carbon isotope value would indicate that marine foods were not likely a major protein source (Mannino et al., 2011a,b). There is one human isotope result from France from this time period, from the site of Abri Faustin, Gironde (Drucker et al., 2005). This human has a d13C value of 18.8‰, and an elevated d15N value of 12.1‰. This site is less than 50 km from the present coastline, and so these values may indicate marine food consumption, but more likely consumption of fish from the Gironde river, including anadromous species such as salmon. A single application of isotope analysis to Late Upper Palaeolithic  (Garciahumans from Spain exists, from the site of Balma Guilanya  et al., 2009). The results indicate, like most of the other Guixe humans from the time period, a broadly terrestrial diet with d13C values of 19.6‰ and 19.9‰ and d15N values of 6.7‰ and 6.8‰. These d15N values are quite low, and similar to the low values reported for the humans from Gough's and Sun Hole Caves in the UK. In both of these studies the associated fauna, especially herbivores, also had low d15N values, likely due to climatic effects (Richards and Hedges, 2003; Hedges et al., 2004) and were interpreted as

Table 1 Isotope data and radiocarbon ages for Late Glacial humans from Europe. Site

Element

Country

14

Gough's Cave Gough's Cave Gough's Cave Gough's Cave Sun Hole 2 Kendrick's Cave Kendrick's Cave Kendrick's Cave Kendrick's Cave San Teodoro 1 San Teodoro 2 San Teodoro 7 Addaura 1 Romito 2 Romito 3 Romito 4 Romito 5 Romito 6 Romito 7 Romito 8 Romito 9 Arene Candide Arene Candide Riparo Tagliente Abri Faustin  Balma Guilanya  Balma Guilanya  Balma Guilanya

Cranium Scapula Cranium Cranium Ulna Humerus Femur Femur Femur Humerus Cranium Cranium Fibula Humerus/Femur Humerus/Femur Humerus/Femur Humerus/Femur Humerus/Femur Humerus/Femur Humerus/Femur Humerus/Femur n.d. n.d. Rib Metacarpal Radius Tooth Cranium

UK UK UK UK UK UK UK UK UK Italy Italy Italy Italy Italy Italy Italy Italy Italy Italy Italy Italy Italy Italy Italy France Spain Spain Spain

12,485 12,565 12,590 12,485 12,415 11,800 11,930 12,090 11,760 12,580 n.d. n.d. 12,890 n.d. n.d. 11,340 10,862 n.d. n.d. n.d. 13,915 10,910 n.d. 13,190 12,370 n.d. 11,095 10,195

C age ± ± ± ± ± ± ± ± ± ±

50 50 50 50 50 90 90 90 90 130

± 60

± 90 ± 70

± 70 ± 90 ± 90 ± 220 ± 195 ± 255

d13C

d15N

Source

19.3 19.0 19.3 19.3 19.8 17.9 18.0 17.7 18.1 20.0 20.0 19.1 19.7 20.0 19.3 19.6 19.7 19.5 19.1 19.5 18.9 20.0 18.9 18.4 18.8 20.0 19.6 19.9

8.5 7.9 7.7 7.6 7.2 13.8 13.4 13.9 13.7 12.5 12.0 11.5 9.6 10.3 10.1 10.0 9.3 8.9 9.7 9.7 12.4 8.9 9.1 13.0 12.1 6.5 6.7 6.8

Jacobi and Higham (2009) Jacobi and Higham (2009) Jacobi and Higham (2009) Jacobi and Higham (2009) Jacobi and Higham (2009) and Richards et al. (2000) Richards et al. (2005) Richards et al. (2005) Richards et al. (2005) Richards et al. (2005) Mannino et al. (2011a) Mannino et al. (2011a) Mannino et al. (2011a) Mannino et al. (2011a) Craig et al. (2010) Craig et al. (2010) Craig et al. (2010) Craig et al. (2010) Craig et al. (2010) Craig et al. (2010) Craig et al. (2010) Craig et al. (2010) Francalacci (1998), Bietti (1987) Francalacci (1998), Bietti (1987) Gazzoni et al. (2013) Drucker et al. (2005)  et al. (2009) Garcia-Guixe  et al. (2009) Garcia-Guixe  et al. (2009) Garcia-Guixe

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indicating diets where terrestrial herbivores were the main source of dietary protein.
3. The Sandalja II site
The Sandalja II site is located about 4 km north-east of the city of Pula on the Istrian peninsula in Croatia (Fig. 1). During mining in the quarry in 1961 a cavern containing Quaternary sediments was
exposed (Sandalja I), while in the following year a second cavern
was found (Sandalja II). Both are, most likely, part of a single, larger
underground complex (Malez, 1979). The name Sandalja I was later used to designate an isolated bone breccia, the faunal remains from which have been attributed to the Early Pleistocene (Malez, 1979; D. Rukavina personal communication). The initial excavations of
the Upper Palaeolithic site of Sandalja were conducted by B. Ba
ci c and were continued by M. Malez, who excavated the site over 22 seasons between 1962 and 1989 (Miracle, 1995). 3.1. Stratigraphy and chronology The deposits at the site are over 8 m thick (Fig. 2), date to the late Upper Pleistocene and have been sub-divided into 9 major levels (Aei) (Malez, 1979, 1990; Miracle, 1995). These levels are composed of compact or sandy clay with large or small rock fragments and contain Upper Palaeolithic (Aurignacian and Epigravettian) industries (Karavani c, 1999, 2003; Karavanic et al., 2013; Malez, 1974, 1979, 1987), fauna (Miracle, 1995, 1996, 2007; Brajkovi c, 2000) as well as human remains (Malez, 1972; Jankovi c et al., 2012). A humic

level (A) containing Bronze Age pottery is the only level known to have been deposited during the Holocene. Levels G, F, and E have previously been radiocarbon dated to between 28 and 23 ka (14C) BP (Srdo
c et al., 1979; Malez and Vogel, 1969). It is likely, however, that problems exist with these conventional radiocarbon dates measured four decades ago, and perhaps over the interpretation of the stratigraphy given that some of the dates are clearly too young for their associated Aurignacian artefacts (Karavani c, 2003; Vogel and Waterbolk, 1972: 133). Because of this we obtained two new radiocarbon measurements on faunal remains collected during Malez's excavations. A sample from level G yielded a date of 10,580 ± 39 BP (OxA-V-2373-48) while a sample from level F yielded a date of 33,355 ± 290 BP (OxAV-2373-49). The later date is in full chronological agreement with the Aurignacian industry from the same level while the former date is too late for the cultural context of level G and attests a degree of stratigraphic mobility. The human remains from level B/s that form the focus of this study were directly radiocarbon dated to 11,025 ± 60 BP (KIA23489) (Table 2), an age similar to conventional radiocarbon dates on charcoal and bone collagen from Layers B/C, B/d, B/s and B/g, as well as an AMS date on charcoal from Layer B/d (Table 3). These dates agree well with the typological attribution of stone tools from Layers B/C to B/g to the Epigravettian industry (Malez, 1974, 1979, 1987; Karavani c, 1999; Karavani c et al., 2013) and thus may be taken as a clear attribution of the human material to the Late Upper Palaeolithic Epigravettian of the region.


Fig. 1. Map of Croatia showing the location of Sandalja.

M.P. Richards et al. / Journal of Archaeological Science 61 (2015) 204e212

207


Fig. 2. Stratigraphy at Sandalja II (after Malez, 1979: Figure 15 and Miracle, 1995: Figure 3.17).

Table 2
Radiocarbon ages of mammal bones (including a possible human left femur fragment) long bones from Sandalja II. Sample no.

Level

Species

14

Lab no.

SA10 SA102 SA103

B/s G F

Mammal (possible Homo sapiens) Mammal (unidentified species) Mammal (unidentified species)

11,025 ± 60 10,580 ± 39 33,355 ± 290

KIA-23489 OxA-V-2373-48 OxA-V-2373-49

3.2. Faunal evidence The mammalian faunal assemblage from Layers B/C to B/g at
Sandalja II is extremely large and diverse, with 8255 identified specimens (NISP) from 30 taxa. The assemblage is dominated by large bovids (%NISP ¼ 33.7, probably all aurochs), followed by badger (%NISP ¼ 12.7), horse (%NISP ¼ 12.4), red deer (% NISP ¼ 10.7), wild boar (%NISP ¼ 7.8), roe deer (%NISP ¼ 7.2), hare (% NISP ¼ 4.1), and fox (%NISP ¼ 3.2, mostly red fox). There is clear evidence from butchery burning marks of the processing of the carcasses of these animals for skins, meat, marrow, and fat. Other larger mammals are present in very small frequencies (% NISP < 1.5) and include the following: giant deer, European elk, ibex, chamois, fallow deer, European ass, marmot, beaver, hedgehog, stoat, weasel, marten, wolverine, wild cat, wolf, lynx, cave lion, brown bear, and cave bear. The complex taphonomy of the assemblage (Miracle, 1995) suggests that some of these remains were collected by non-human predators (e.g. hedgehog, stoat, weasel, marten) and/or derived from episodes of cave use by animals (e.g. marmot, wolf, lynx, cave lion, brown bear, cave bear); in

Table 3
Radiocarbon ages of the upper stratigraphic layers at Sandalja II. Layer

Age

B/g B/s B/d B/d B/C

10,830 12,320 10,990 10,140 13,050

± ± ± ± ±

50 100 60 160 220

Laboratory Number

Reference

Grn-4976 GrN-4978 CAMS-12062 Z-2421 Z-2423

Malez and Vogel (1969) Malez and Vogel (1969) Miracle (1995) Obeli c et al. (1994) Obeli c et al. (1994)

C age

both cases these rare taxa would not have contributed to the human diet. A further problem arises from the relatively coarse excavation methods that were used, and the likelihood that some remains may be out of stratigraphic context (e.g. cave bear).
Sandalja II contains relatively rich fish, amphibian, and reptile assemblages, which have been identified and analysed by Paunovic (1984, 1987). The excavated sediment was not systematically sieved, however, and we suspect that there has been a strong bias against the recovery of remains of fish and other smaller animals. Out of a total of 1533 remains identified to one of these classes from layers C/d to B/g, 645 (42%) were determinable to genus and/or species (Paunovi c, 1984:43). Over half of this identified assemblage (%NISP ¼ 56) derives from seven species of amphibian (green toad predominates with 29.7% of NISP). A total of 158 remains (% NISP ¼ 24.5) were identified to ten fish species, of which the common pike was most frequent (%NISP ¼ 12). Other fish species are relatively rare (%NISP < 2.5) and include rudd, common carp, tench, crucian carp, bream, perch, barbell, dace, and N€ aling. Only freshwater fish have been identified in the assemblage, and of these species most prefer the slower moving waters of larger rivers, ponds, or lakes (Paunovi c, 1984: 35). Reptiles from eight species account for 19% of the amphibian/fish/reptile assemblage; European pond turtle is the best represented taxon (%NISP ¼ 5.9). A large assemblage (NISP ¼ 911) of avifaunal remains was studied by Vesna Malez (Malez-Ba
cic, 1979), and is currently being revised by Ankica Oros-Sr
sen (Oros-Sr
sen et al., 2014). A number of bird bones show evidence of burning and butchery, although the taphonomy of this assemblage also appears to be complex (OrosSr
sen et al., 2014). Finally, a number of microfaunal remains including small rodents, bats, and insectivores were also recovered.

208

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This assemblage has not been studied; our working hypothesis is that these were introduced into the deposits by roosting birds and other non-human predators. The taphonomy of the amphibian/fish/reptile assemblage has not been studied. Some of these remains presumably derive from animals that died of natural causes in the cave (e.g. the amphibians, lizards, and snakes). Fish, lizards, snakes, and amphibians also could have been brought to the site by roosting birds and/or other non-human predators. We suspect, however, that turtle and fish were most likely consumed by humans.
In summary, the vertebrate assemblages from Sandalja II indicate regular and systematic consumption of large, terrestrial mammals by Late Upper Palaeolithic humans. In addition to these some fish, birds, and turtle were also consumed. Given the numerical dominance of larger mammal remains as well as their much larger body size, one could infer from these assemblages that terrestrial mammals were the primarily source of terrestrial protein in the Late Upper Palaeolithic human diet. 4. Methods We sampled between 100 and 200 mg of bone from three hu
man and 29 faunal samples from the Sandalja II site (Table 4). Collagen was extracted following a modified Longin (1971) method (Richards and Hedges, 1999) with an ultrafiltration step (Brown et al., 1988). Carbon and nitrogen isotope measurements were undertaken at the Department of Archaeological Science, University of Bradford, U.K., and the Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany. The isotope data, and the collagen preservation criteria are presented in Table 4. Only samples which have acceptable C:N ratios (2.9e3.6, DeNiro, 1985) are included here.

There are at least two archaeological phases represented at the site. The older phase (levels F and G) dates to the Aurignacian period, ca. 33,000 BP and the other younger phase (levels B/CeB/g) dates to the late Pleistocene Epigravettian ca. 11,000 uncal BP. The humans and all of fauna that we sampled (excepting two unidentified long bone shafts submitted for radiometric dates) belong to the younger phase. In order to interpret the human isotope values it is necessary to compare them to isotope values of contemporary fauna, as fluctuations in carbon and nitrogen isotope values are observable in Europe throughout the Pleistocene, which are most likely linked to climate change (Richards and Hedges, 2003). Level B is subdivided into upper (B/g), middle (B/s) and lower (B/d) levels, wit the majority of fauna and all human remains deriving from the middle level (B/s) of layer B. We do not discuss further the isotope results from the two samples from the lower levels F and G. Of the three human remains we have two clearly identified human bone fragments (14015 [S-EVA-5256], 14021 [S-EVA-5277]) and one (14052 [SA10]) that is too fragmentary to definitively identified as human, so we refer to it as a ‘possible’ human in Tables 2 and 4 below. We have labelled this sample as ‘human’ in Figs. 3 and 4. We have plotted first the isotope values of humans and fauna from the same middle section of layer B (B/s) in Fig. 3, and the average values of humans and fauna from all of the sections of layer B in Fig. 4. 5. Results As can be seen in Figs. 3 and 4, and Table 4, the human carbon and nitrogen isotope values do not reveal any measurable consumption of marine foods. Their nitrogen isotope value are in fact 3e4‰ higher than those of the carnivores we measured. With

Table 4
Isotope data for humans and fauna from Sandalja II. Lab no.

Submitter no.

Species

Level

d13C

d15N

%C

%N

C:N

%Collagen (>30 kDa)

S-EVA-5252 S-EVA-5253 S-EVA-5254 S-EVA-5255 S-EVA-5256 S-EVA-5257 SA10

SDL-24 SDL-22 SDL-25 SDL-23 SDL-11 SDL-12 n/a

B/d B/d B/s B/d B/s B/s B/s

24.1 22.6 22.7 23.7 20.6 20.6 20.8

9.1 9.2 9.3 8.5 13.6 14.0 13.1

41.0 36.3 32.9 30.7 40.9 35.8 44.2

13.7 12.2 11.2 10.6 14.2 11.4 14.9

3.5 3.5 3.4 3.4 3.4 3.6 3.5

0.6 1.0 0.3 0.8 0.8 0.3 2.4

S-EVA-5258 S-EVA-5259 S-EVA-5262 S-EVA-5260 S-EVA-5263 SA6 S-EVA-5264 S-EVA-5271 SA7 S-EVA-5265 S-EVA-5266 S-EVA-5267 SA4 S-EVA-5268 S-EVA-5269 S-EVA-5270 SA5 S-EVA-5273 S-EVA-5274 S-EVA-5275 S-EVA-5276 SA8 SA100 SA102 SA103

SDL-18 SDL-19 SDL-21 SDL-13 SDL-14 n/a SDL-16 SDL-17 Sandalja SDL-10 SDL-9 SDL-8 Sandalja SDL-7 SDL-5 SDL-6 Sandalja SDL-4 SDL-1 SDL-2 SDL-3 Sandalja Sandalja Sandalja Sandalja

Pike 567 Pike 1126 Pike 1327 Pike 1390 Human 14015 Human 14021 Possible human left femur fragment 14052 B/S Felis 11001 Felis 11007 Felis 11013 Canis 11110 Canis 11089 Canis lupus B/s Panthera 11893 Panthera 3570 Panthera spelaea Bos 2363 Bos 2364 Bos 2367 Bos primigenius prox. humerus Cervus 2381 Cervus 2387 Cervus 2408 C. elaphus proximal humerus Equus 5249 Equus 5257 Equus 5309 Equus 5338 Ursus spelaea molar root B/S Green teach ? Long bone mammal ? Long bone mammal

B/C B/C B/C B/C B/g B/s B/C B/s B/s B/s B/s B/s B/s B/s B/s B/s B/s B/s B/s B/s B/s B/s ? G F

21.1 19.5 20.3 18.6 18.5 20.2 18.5 18.1 19.2 20.5 19.7 19.8 21.0 20.4 21.2 21.1 20.4 20.4 20.4 20.5 20.4 21.7 19.8 20.2 18.9

11.4 8.6 6.5 9.1 12.1 9.5 9.7 10.2 9.2 8.0 7.8 6.0 6.4 4.6 4.1 4.3 2.7 7.3 8.4 6.7 4.9 6.9 8.0 8.4 8.0

40.9 41.0 40.6 35.3 32.5 29.8 38.4 41.2 42.2 41.2 42.1 41.9 42.1 41.6 39.6 23.3 37.5 40.1 40.4 40.7 40.6 37.8 43.6 41.1 25.9

14.6 14.7 14.3 12.6 11.6 9.9 14.0 15.1 14.8 14.8 15.1 15.0 13.82 14.7 13.9 8.1 12.3 14.1 14.7 14.5 14.3 12.8 14.3 13.9 8.5

3.3 3.3 3.3 3.3 3.3 3.5 3.2 3.2 3.3 3.3 3.3 3.3 3.6 3.3 3.3 3.4 3.6 3.3 3.2 3.3 3.3 3.5 3.6 3.5 3.6

1.4 2.0 0.6 1.5 0.8 1.4 3.2 1.5 1.2 1.6 2.1 1.2 0.6 0.7 0.4 0.7 0.3 0.3 1.4 0.5 0.6 0.2 0.4 2.0 0.7

M.P. Richards et al. / Journal of Archaeological Science 61 (2015) 204e212

Fig. 3. Bone collagen carbon and nitrogen isotope values of humans and fauna from sub level ‘B-middle’ (B/s).

nitrogen isotope analysis consumers will generally be higher by 3e5‰ than the nitrogen isotope values of their protein sources (i.e. the protein they consume: Schoeninger and DeNiro, 1984; Hedges
and Reynard, 2007). It is clear that the Sandalja II carnivores have nitrogen isotope values that are 3e5‰ higher than those of the site's cervids, bovids, and equids, confirming the archaeozoological conclusion that these herbivores were likely to have been their main protein source. The human d15N values are much higher than this, however, and one cannot therefore explain their values in terms of a diet in which protein derived mainly from herbivores. Likewise, although there is faunal evidence for the systematic consumption of badgers and carnivores like wild cat and fox, which would result in elevated human nitrogen isotope values, these species would have been in small quantities relative to the other fauna at the site, and so it is unlikely that they would have formed the main protein source in human diets. We therefore interpret the human high d15N values as reflecting the consumption of protein from a different ecosystem, namely freshwater. The isotope values
of pike from Sandalja II (average d15N value of 9‰), which could indicate that this taxon formed the main protein source for the humans, who possess values 3e5‰ higher than these freshwater fish. With isotope studies such as these we are necessarily limited to building an isotope food web using the fauna that is available to us

Fig. 4. Bone collagen carbon and nitrogen isotope values of humans and fauna from level B/s.

209

from the site; plant remains rarely, if ever survive, thus are generally invisible in these studies. As a proxy for these we used the herbivore values, as we assume that they are 3e5‰ higher than the plants they consumed, thus we may assume that the plants in Late
Upper Palaeolithic Sandalja II had nitrogen values of 3e5‰, which is what we would predict for temperate Europe at this time, based on numerous studies of modern plants and animals from similar environments. If plant protein was a major component of human diets in the region during the Late Glacial we would expect humans to have d15N values much closer to those of the herbivores; as they are much higher we conclude that plants were not a major source of dietary protein, although, of course, they could have constituted (and likely did) an important source of carbohydrates and micronutrients. As a further isotopic indicator of freshwater fish consumption we attempted to undertake sulphur isotope analysis (Hu et al., 2009) of these humans, but were unfortunately not able to obtain enough collagen for this analysis.
We therefore interpret the Sandalja II human isotope values as indicating that these Late Upper Palaeolithic humans derived the majority of their dietary protein over their lives from freshwater fish. This stands in striking contrast to the zooarchaeological evidence from the site. 6. The wider isotopic context To compare our results with existing isotope values of European Late Upper Palaeolithic humans and to explore the possibility of trends through time, we have plotted the carbon and nitrogen isotope values of humans that have been directly radiocarbon dated in Fig. 5. As can be seen, most of the humans have d13C values clustering around 20‰, which is the usual value for a diet where the protein was largely derived from terrestrial sources. The two clear exceptions are the humans from Kendrick's Cave, Wales and the humans from Romito, Riparo Tagliente and Abri Faustin where marine protein was an important e although not dominant e part of their diets. The d15N values are much more variable, although again most d15N values are between 9 and 10‰, indicating that terrestrial animals were the main protein source of the humans sampled. The lowest d15N values (ca. 7e8‰) are from Gough's and Sun Hole  in Spain, which in each case Caves in England and Balma Guilanya can be explained in terms of diachronically fluctuating d15N values rather than specifically dietary terms. As discussed above, at Gough's and Sun Hole all of the associated non-human fauna have lowered d15N values, a pattern observed for fauna across Western Europe around this time (Richards and Hedges, 2003; Hedges et al., 2004; Drucker et al., 2003, 2012; Stevens and Hedges, 2004). The  humans are also associated with herbivores with Balma Guilanya low d15N values (in both sites, the average herbivore value is ca. 1.7‰). Interestingly this phenomenon of lower d15N values at this time period in Western Europe does not seem to occur in the Italian Late Upper Palaeolithic sites, however, observed, for example among the fauna of Paglicci Cave (Iacumin et al., 1997). The Kendrick's Cave, Romito, Riparo Tagliente and Abri Faustin individuals have higher d15N values than the samples from elsewhere, likely due to the consumption of marine protein. The other
two individuals with high d15N values are Sandalja II and San
Teodoro. San Teodoro, like Sandalja II, possesses terrestrial-like d13C values but very elevated d15N values, most likely indicating fresh
water fish consumption. Although at San Teodoro, unlike Sandalja, some of the herbivores, especially aurochs, had relatively elevated d15N values, leading Mannino et al. (2011a,b) to conclude that the human d15N value of 12.5‰ likely indicated a diet characterised by a mix of terrestrial game and freshwater fish, the latter probably brown trout (Salmo trutta). As this was apparently not the case at

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Fig. 5. Bone collagen carbon and nitrogen isotope date for directly radiocarbon dated Late Upper Palaeolithic humans from Europe.


Sandalja II, we conclude that the elevated d15N values indicate that freshwater fish was likely the main source of dietary protein for the humans sampled from the site. 7. Discussion
The Epigravettian hunteregatherers of Sandalja II were clearly repeated and adept hunters of large herbivores such as aurochs, horse, red deer and a number of smaller taxa. Our results demonstrate that their small animal procurement extended into the freshwater realm, and that the procurement of fish formed a major component of their protein intake. This forms a clear example of the dietary importance of small animals, in the context of a diverse hunted and (one assumes) trapped faunal spectrum on land and on water. It serves as a useful warning that while direct reading of faunal assemblages provides useful indications of the relative importance of individual taxa among large herbivores, this does not necessarily translate into the relative importance of large herbivores among the diet as a whole. As Owen (2002) has noted, archaeologists have paid little attention to small animal use in Late Upper Palaeolithic diets. The few studies that now exist e those discussed above and our own results e demonstrate that the contribution of small animals to diet in the Late Glacial could be significant, and should therefore not be written off as simply

ancillary to the more important tasks of hunting game. As additional studies emerge we will no doubt be able to nuance our understanding of Late Glacial diet further, but for now we note that several research questions emerge. It is easy to understand why terrestrial herbivores dominate SE European Late Glacial faunal assemblages, notably Klithi, Greece (ibex: Gamble, 1997), Boïla, Greece (chamois and ibex: Kotjabopoulou et al., 1999), Badanj, Herzegovina (red deer: Miracle and Sturdy, 1991; Miracle, 1995, 1996; Whallon, 1999), Kopa
cina, Croatia (red deer: Miracle, 1995, 1996), Pupi cina, Croatia (red deer: Miracle, 2007), Ve
sanska, Croatia (red deer: Miracle, 2007), Vela spila, Croatia (red deer: Spry-Marques, 2012), and Treba
cki Kr
s, Montenegro (red deer, ibex and chamois: Dimitrijevic., 1999). Despite this, evidence for Late Glacial small animal use is known in southern Europe, e.g. Grotta di Polesini and Grotta dei Cervi, Italy (pond tortoise, duck, fish: Mussi, 2001). Second is the question as to whether humans avoided any significant consumption of locally available plant foods. Humans were apparently absent from a florally diverse environment at Ezero in SE Europe, for example (Magyari et al., 2013). This might seem strange in the context of a growing broadening of the dietary spectrum (e.g. Stiner, 2001): why, if humans were clearly capable of exploiting small resources in a variety of ecosystems might they have ignored plants or not exploited areas in which they were

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diverse? Should one be surprised at this, or does it simply reflect the vagaries of preservation and recovery? Plant remains have certainly been under-represented and under-sampled in cave sites in particular (Bailey, 1997), although some evidence exists, e.g. the remains of edible plant remains in Late Glacial contexts at Arene Candide, Italy (Cardini, 1946). A third question is whether one can assume that individual isotopic diets are representative of the diet of conspecifics (i.e. that diets were homogeneous) or whether one might expect individual dietary diversity in this period, and if so to what extent. At Romito, for example, isotopes suggested that the proteinacious element of the diet of all individuals analysed bar one was dominated by terrestrial herbivores but that one individual possessed a significant marine component (Craig et al., 2010). 8. Conclusions We applied isotope analysis to human and faunal remains from
the Late Upper Palaeolithic levels at Sandalja II in Croatia. We found that the isotope data from the fauna that was available for analysis and contemporary with the humans that these humans likely obtained the majority of their protein from freshwater fish, which contrasts with the site's zooarchaeological evidence for the consumption of large amounts of meat, marrow, and fat from large terrestrial mammals in the same level. Our results add to the growing evidence of increasingly intensive use of aquatic resources in the European Late Upper Palaeolithic (Gazzoni et al., 2013; Drucker et al., 2005; Mannino et al., 2011a,b), a trend we see continuing into the Mesolithic and culminating with humans depending almost entirely on marine resources in coastal northwest Europe in the Late Mesolithic period (e.g. Richards et al., 2003a,b; Fischer et al., 2007). We hope that isotope research in the Late Glacial will continue and expand to other areas of Europe, so we can build up a more comprehensive picture of Late Upper Palaeolithic dietary adaptations and especially the increasing intensification and diversification of resources e especially aquatic e that continues into the Holocene. Acknowledgements We would like thank to the late Maja Paunovic for her help in initiating this project and in giving us permission for sampling. We would also like to thank Gordana Jambre
si c for her help in selecting the samples at Institute for Quaternary paleontology and geology of the Croatian Academy of Sciences and Arts initially, and to Dejana Brajkovi c for permission to sample additional materials. This research was funded by the Natural Science and Engineering Research Council (NSERC) of Canada and the Max Planck Society. References Bailey, G.N. (Ed.), 1997. Klithi: Palaeolithic Settlement and Quaternary Landscapes in Northwest Greece. Cambridge. McDonald Institute Monographs. Bietti, A., 1987. Some remarks on the new radiocarbon dates from the Arene Candide Cave (Savona, Italy). Hum. Evol. 2, 185e190.
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