Ice Age apparel – changing prey patterns towards the Last Glacial Maximum and the role of reindeer fur for clothing at Kammern-Grubgraben

doi:10.21203/rs.3.rs-4129631/v1

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Ice Age apparel – changing prey patterns towards the Last Glacial Maximum and the role of reindeer fur for clothing at Kammern-Grubgraben

https://doi.org/10.21203/rs.3.rs-4129631/v1

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Isotope analyses on bone collagen from hunted animals at the Upper Palaeolithic sites Krems-Hundssteig, Krems-Wachtberg and Langenlois A (33 − 29 ka), and Kammern-Grubgraben (23 ka) indicate a strong reduction of ecological herbivore niches towards the Last Glacial Maximum. This change is accompanied by a clear shift in human prey pattern from a more diverse and balanced spectrum of hunted species in the earlier period to a clear focus on reindeer at Kammern-Grubgraben, particularly on young individuals. Moreover, new analyses of the osteological material show that the site with its massive stone constructions and high amount of lithic and faunal material was occupied exclusively in winter. In this paper, we present new results on the age and sex distribution of reindeer at Kammern-Grubgraben. We argue that winter-hunting of young reindeer, in addition to its role in providing energy-rich nutrition and raw material for organic tools, is also indicative of a focus on obtaining high-quality raw material for clothing. The fur of young reindeer in winter is particularly valuable and convenient for the production of clothing for cold environments. Together with the recovered large number of eyed needles, a tool for tight and regular seams, our findings suggest that the production of clothing and other goods made of fur and skin was an important activity at Kammern-Grubgraben.

Upper Palaeolithic

Lower Austria

Gravettian

Epigravettian

subsistence change

reindeer fur use

This study presents changes in the use of recourses between MIS 3 and MIS 2 sites in Lower Austria (Fig. 1), as revealed by osteoarchaeological and archaeological evidence and by environmental changes based on isotopes of animal bones. The analysis is based on the archaeological sites Krems-Wachtberg (excavation periods of 1930 and 2005–2015), Krems-Hundsteig (excavation period 2000–2002), Langenlois A and Kammern-Grubgraben.

The Upper Palaeolithic (UP) site cluster at Krems includes, among others, the sites Krems-Wachtberg 1930, Krems-Wachtberg 2005–2015 and Krems-Hundssteig 2000–2002 which are considered in this study. All three sites are multi-layered and include both occupations with evident features such as hearths and burials, and find layers representing palimpsest accumulations created by natural slope processes. At Krems-Wachtberg 1930 (excavated in 1930) the finds from these different contexts were not separated (Einwögerer, 2000). Seasonal conclusions in case of more than one season represented must therefore be interpreted with caution. At Krems-Hundssteig, the bulk of the faunal remains derives from layers HU-AH 3.21, HU-AH 3.23 and HU-AH 3.24 (Neugebauer-Maresch, 2008; Fladerer & Salcher 2010) which represent palimpsests of finds from different occupations accumulated by slope movements. A similarly insecure context must be attested for layers HU-AH 3.3 and HU-AH 3.4. This leaves only HU-AH 3.22 with hearth D as an archaeological context of sufficient integrity. The other secure archaeological contexts at Krems-Hundssteig 2000–2002 did unfortunately not provide (diagnostic) faunal remains and are therefore not relevant for this study.

At Krems-Wachtberg 2005–2015, layer WA-AH 4.4/4.3 represents the remains of a distinct occupation with evident structures such as hearths, pits, and burials, and therefore qualifies for seasonal conclusions for this occupational event. The faunal material is often highly fragmented and burnt by human activities. On the other hand, layer WA-AH 4.11 is a palimpsest comparable to layers HU-AH 3.2 (layers HU-AH 3.21, HU-AH 3.23, and HU-AH 3.24) at Krems-Hundssteig (Händel 2017). This is supported by archaeological, sedimentary and radiocarbon data (Händel et al., 2021b). Based on archaeological and chronological data, Händel (2017) proposes three phases of Gravettian occupation at the UP-site cluster of Krems, while each of these phases is characterised by several distinct presences of varying duration.

Langenlois A is a single-layer site and is interpreted as representing one single occupational context with evident settlement structures such as hearths, pits, and distinct activity areas (Einwögerer, 2019). Seasonal conclusions can therefore be considered indicative for the occupation connected to these features.

The LGM site Kammern-Grubgraben shows two main archaeological layers, KG-AH 1 and KG-AH 2, both of which show a great variety of human activities. The materials most probably accumulated in the course of repeated visits. While layer KG-AH 2 shows a high number of evident anthropogenic features and structures including hearths and a variety of constructions made of stone slabs carried into the site with no signs of substantial post-occupational mixing caused by slope processes, such disturbances seem likely to have affected layer KG-AH 1 (Händel et al. 2021a). Seasonality results are therefore only meaningful for the occupation phases associated with the evident structures in the KG-AH 2 layer. In case of more than one season represented, results for layer KG-AH 1, however, should be treated with caution.

Isotopic analyses of bone collagen of the hunted fauna were recently published for the main find layers at the UP sites Krems-Wachtberg 1930, and Krems-Hundssteig 2000–2002 (33 − 31 ka cal BP), as well as at Langenlois A (30.8–29.2 ka cal BP), and at Kammern-Grubgraben (24 − 20 ka cal BP). These results show a strong reduction of ecological herbivore niches towards the Last Glacial Maximum (Reiss et al., 2023). The main UP occupations at the Krems-Wachtberg sites (Einwögerer, 2000; Einwögerer et al., 2009; Händel, 2017), at Krems-Hundssteig 2000–2002 (Neugebauer-Maresch 2008; Händel 2017) and at Langenlois A (Einwögerer, 2009, 2019) fall into the most recent stage of Marine Isotope Stage (MIS) 3 and thus pre-date the LGM (Clark et al., 2009; Mix et al., 2001). In contrast, the studied UP occupations at Kammern-Grubgraben (Haesaerts, 1990; Haesaerts et al., 2016; Händel et al., 2021a; Lengyel et al., 2021; Montet-White, 1990, 1994) are situated within MIS 2, and thus the LGM.

From the older period (33 − 29 ka cal BP) to the younger period (24 − 20 ka cal BP), climatic conditions were characterised by constantly declining insolation and temperature, interrupted only by short interstadials (Andersen et al. 2004; Kindler et al. 2014). The radiocarbon age ranges of the archaeological contexts considered in this study, Krems-Wachtberg 1930 and Krems-Wachtberg 2005–2015 (layers WA-AH 4.11, WA-AH 4.4) and Krems-Hundssteig 2000–2002 (layers HU-AH 3.2, HU-AH 3.3/3.4) cover Greenland Interstadial (GI) 5.2 and Greenland Stadials (GS) 6 and 5.2, the range of Langenlois A covers GI 5.1 and GS 5.1, and the one of Kammern-Grubgraben (layers KG-AH 1, KG-AH 2) covers GI 2.2 and 2.1 and GS 3, 2.2 and 2.1 (Rasmussen et al., 2014). Based on the radiocarbon ages, it is therefore currently not possible to say with certainty whether any of the referred occupations took place during suggested stadial conditions (Reiss et al., 2024).

The faunal remains from Krems-Wachtberg 1930, Krems-Wachtberg 2005–2015 and Krems-Hundssteig 2000–2002 were already well documented previously (Fladerer, 2001; Fladerer & Salcher-Jedrasiak, 2010; Fladerer et al., 2014; Händel et al., 2015; Händel, 2017), while the osteological material from Langenlois A and Kammern-Grubgraben (Brandtner/Klima excavation) is published here for the first time.

All faunal remains of the material from Langenlois A were recorded for the present analysis. However, osteological finds from the archaeological excavations at Kammern-Grubgraben of the 1980s and 1990s by A. Montet-White/F. Brandtner and by F. Brandtner/B. Klima, provide the main material basis for this study. The finds from the Montet-White/Brandtner excavation at Kammern-Grubgraben have already been presented in several publications (Montet-White, 1988, 1990, 1994; West, 1997; Williams, 1998), but were re-analysed together with the hitherto unrecorded finds from the excavation by Brandtner and Klima using the same protocol and standards. Because of the large quantity of material (354 large standard boxes), we opted for a selective method when analysing the osteological material. Only finds that can be assigned unambiguously to an archaeological horizon were chosen. The nomenclature for the layers changed during research: Montet-White/Brandner assigned archaeological layers (AL) 1 to 4, and Brandtner/Klima cultural layers (KS) 1 to 4. Recent excavations conducted by the Austrian Academy of Sciences, however, can only confirm two major archaeological units for this sequence: only the uppermost layer, AL resp. KS 1, is clearly separable due to its position in relation to the other layers, and was renamed KG-AH 1, whereas AL resp. KS 2–4 are neither defined by individual sedimentary properties, nor separated by archaeologically sterile sediments, and are therefore considered as a single unit, KG-AH 2 (Händel et al., 2021a). It should be noted that KG-AH 2 allows for distinguishing a high variety of activities – this spatial patterning is however not considered in our study. Here, we follow the nomenclature introduced by Händel et al. (2021a) and consider the two main archaeological units KG-AH 1 and KG-AH 2. In horses and reindeer, all postcranial measurable fragments and those with age characteristics, all teeth and, in reindeer, the ossa frontalia with and without antler remains were recorded. For the other species, all remains were recorded. This paper presents the documented data, but detailed statements on KG-AH 1 will be the subject of future analyses.

Animal species could not always be determined at species level, since individual genera cannot be reliably separated on a morphological basis, e.g. in the case of the Arctic and the European hare. We have therefore decided to also use the term "genus" in addition to the term "species".

For the quantification of the identified animal specimens two quantification methods are provided. The number of identified specimens (NISP; Payne, 1975) is used which defines the maximum number of taxa represented (Klein & Cruz-Uribe 1984, 30). The minimum number of individuals (MNI) was calculated on basis of the most frequent skeletal element, in combination with the bone part, the body side, age and sex (Reitz & Wing, 1999, 195). Each quantification method has advantages and disadvantages and is discussed intensively such as the MNI calculation which involves some uncertainties and is not necessarily identical to the actual number of hunted animals (see summary in Reitz & Wing, 1999, 194–202). However, the MNI is one of the most common calculations in Palaeolithic archaeology and thus allows the use of most comparisons. At the very least, the MNI gives an approximate value of the relative importance of the different species and genera and allows a comparison between the two periods under study.

[header 2] Dental cementum analysis

Thin sections of teeth were prepared to determine the seasonality of adult reindeer and other species. Previous analyses on seasonality on annual dental cementum rings on the material from Kammern-Grubgraben were not successful due to patchy cementum structure in the selected specimens (Whitney, 1992, 40; Williams, 1998, 88). Therefore, we tried two different methods to achieve better results. Initially, we tried to decalcify the teeth (Suppl. 1) but because of the fragility of the osteological material this method did not prove successful. Embedding teeth in resin worked better, especially for teeth still embedded in the jawbone.

[header 2] Odontochronology and tooth wear of reindeer

Odontochronology, i.e., the determination of tooth age based on the eruption sequence of teeth, and tooth wear was used to determine biological age and season of death. The knowledge of tooth development and tooth eruption in the reindeer is based on knowledge of tooth development in other Rangifer populations (Table 1).

Table 1

Eruption and tooth wear of the fourth deciduous premolar (Pd4), and the permanent molars (first molar M1, second molar M2, and third molar M3) according to Brommé-Skunke (1952) and Pasda (2009, Table 8)
biological age	tooth type and tooth wear
birth (~ end of May/beginning of June as is known from recent populations, however calving time varies)	Pd4 erupting shortly after birth
3 months	Pd4 in occlusion, little wear
3–5 months	M1 erupting, Pd4 moderately worn
6 months	alveolus of M2 starts opening, M1 slight wear
7–13 months	M2 erupting, alveolus of M3 starts opening
13–18 months	M3 erupting, M2 slight wear, Pd4 heavily worn

For the assessment of individual age of reindeer, the tooth wear stages of Pd4, M1, M2, and M3 were initially recorded for each tooth according to the wear scheme developed by Van den Berg et al. (2021, absolute scheme/Fig. 4). Alphabetical labels of this scheme were replaced by numerals, starting with 1 for a. The information was supplemented with the wear stages "-1" for "in breakthrough" and "0" for "not worn". In addition, the greatest width and the greatest thickness of the tooth crown were measured. During the analysis, however, it emerged that the application of this method developed on Svalbard reindeer (Rangifer tarandus platyrhynchus) was not suitable on the archaeological material of Kammern-Grubgraben as several issues affect the utility. First, the assessment of the degree of wear in the reindeer teeth from Kammern-Grubgraben was not always unambiguously possible. Furthermore, the assessment of several teeth in one jaw showed that the degree of wear varied between individual teeth within a jaw (Suppl. 2). Finally, the combination between the tooth wear stages of the individual teeth under study shows a large variation. It was therefore not possible to determine a precise biological age with this method. Therefore, the determination of age and season of death was done with a combination of tooth wear stage and tooth dimensions on the 4th milk premolar (Pd4), as it is only present in calves up to the second winter of life and the start of development and lifespan of the tooth can be narrowed down more precisely than is possible with other teeth. The applied method was first developed and tested on a Greenlandic caribou population, where the age of the young animals could be defined relatively well (Pasda, 2009). Based on the Greenland caribou a wear scheme was created for the lower and upper jaws (Fig. 2). The scheme starts with Pd4’s wear stage 3 (as defined by Van den Berg et al., 2021, Fig. 4), since there was no tooth with a wear stage less than 3 available in the material of Kammern-Grubgraben. As tooth eruption and development of M1 and M2 are also relatively well defined, the assessment of tooth wear was also applied to these teeth. The variation of the eruption in the M3 is quite large, so that this tooth was not included. The combination of tooth crown wear stage tooth crown dimensions of the Pd4 is shown in Table 2.

Table 2

Tooth dimensions, wear stage, and age characteristics at the 4th milk premolar (Pd4) on caribou from Central West Greenland (Pasda, 2009), “M” mean value of the respective wear stage
tooth crown length	tooth crown height	wear stage	age class	age in months	season
		Maxilla
16.64	9.93	-1	fetal/shortly before birth		spring
17.52	6.55	5	juvenile	6–9	winter
16.41	5.90	5	juvenile	6–9	winter
16.08	5.42	5	juvenile	6–9	winter
16.13	2.51	7	subadult	9–12	winter
15.43	2.51	7	subadult	9–12	winter
16.25	5.60	M 5
15.78	2.51	M 7
		Mandibula
22.51	10.08	-1	fetal/shortly before birth	0	spring
21.32	9.19	-1	fetal/shortly before birth	0	spring
22.19	10.07	-1	fetal/shortly before birth	0	spring
22.31	6.29	4–5	juvenile	6–9	winter
21.21	5.04	5	juvenile	6–9	winter
23.02	5.41	5	juvenile	6–9	winter
23.23	6.38	5	juvenile	6–9	winter
21.66	5.55	6	juvenile	6–9	winter
21.25	4.19	6	juvenile	6–9	winter
21.18	3.88	7	subadult	9–12	winter
20.47	4.12	7	subadult	9–12	winter
21.48	5.35	7	subadult	9–12	winter
20.69	2.57	7	subadult	9–12	winter
22.01	9.78	M fetal
22.44	5.78	M 5
20.96	3.98	M 6–7

Figure 3 shows tooth crown dimensions and wear stages of the Pd4 according to upper and lower jaw in the West Greenland caribou, ranging from fetal individuals up to animals that died in the first and second winter of life (Pasda, 2009). Calves that died between birth and winter, were not available. While two distinct groups of animals that died during the first and second winter are observable in the data from the lower jaw, the upper jaw shows a certain overlap in the values. However, the mean values of the data are clearly separated from each other. The first data with wear stage 5 come from animals that were about 6–9 months old, i.e., that died between December and March. Wear stages 6–7 represent animals that died in the second winter of life but cannot be broken down more precisely. The temporal dimension in relation between tooth dimensions and wear stage is clearly visible in the development of the data in the diagram from top right to bottom left.

[header 2] Sex determination of adult reindeer

The sex of reindeer can be clearly determined by the pelvis (Pasda, 2009, 11, Fig. 8). In KG-AH 2, there are eight fragments with the relevant diagnostic areas. Of these, three were identified as female and two as male. No pelvis with the corresponding diagnostic section was available for KG-AH 1. As this number is generally low in view of the large amount of osteological material, sex determination was also done via bone size. Since reindeer and caribou have a clear sexual dimorphism, the dimensions are a potential means of determining the sex of individual skeletal elements. Since the dimensions for female and male reindeer of this time and area are not known, data from a recent Rangifer population was used. However, populations differ in their absolute sizes (Meldgaard, 1986, 15), and possibly also in their proportions. However, as the archaeological material from Kammern-Grubgraben was heavily fragmented, there was not a single complete long bone available that would have allowed to determine the height at withers, so that only isolated joint ends could be used. Nevertheless, width and length measurements of the joint ends with fused epiphyses can reveal sexual differentiation.

To determine the sex, data from a Greenlandic caribou population was used, in which skeletal elements of sexually known animals was measured. The application was first tested on the distal tibia (data see Pasda, 2009, Table 68). The sexual dimorphism in the Greenland caribou is clearly visible, but there is also a certain area of overlap (Fig. 4). In addition, measurements of a late glacial assemblage of female reindeer from Stellmoor in North Germany were used (data see Pasda, 2009, Table 416).

The data of the female animals from Stellmoor lie within the variation range of the female Greenland caribou and confirm a similar variation in the dimensions of the female reindeer from North Germany as that of the female caribou of Greenland. The variation in the data from Kammern-Grubgraben is relatively large and scatters over the entire range of the Greenlandic comparative data. However, the comparison shows certain differences in proportions. The tibiae appear to be similar in width, while certain differences can be recognized in the thickness. The tibiae of the female reindeer from Stellmoor are slightly thicker than those of the Greenlandic females. This trend is also visible in the larger dimensions of Kammern-Grubgraben compared to the male Greenlandic caribou. At least in terms of their width, the reindeer tibiae from Kammern-Grubgraben appear to have values comparable to those of the Greenland caribou, and discrimination with these dimensions seems thus possible.

Because of the differences in proportions noted in tibia thickness, only the width was considered for determining the sex of adult animals. As only skeletal components with epiphyseal fusion happening after 18 months of age were utilised (epiphyseal fusion according to Pasda, 2009, Table 13, definition of measurements see Pasda, 2009, 14–15), the data was significantly constrained, since only radius distal, tibia distal and calcaneus offered a sufficiently large number of measurements (Suppl. 3, sex determination based on data in Pasda, 2018).

[header 2] MNI calculation for Kammern-Grubgraben

The MNI of the represented female, male and indeterminable reindeer was calculated separately for the archaeological horizons (AH) (Table 3).

Table 3

Kammern-Grubgraben. Determination of MNI by sex in postcranial skeletal elements of reindeer over 18 months of age. The grey shaded cells show the MNI of the identified male, female, or non-sexed adult animals. Data see Supplement 3.
skeletal element and dimension	female sinistral	female dextral	male sinistral	male dextral	? sinistral	? dextral
			KG-AH 2
Radius Bd	1	2	4	3	6	11
Tibia Bd	11	5	3	4	1	6
Calcaneus GL	5	3	1	2	3	3
			KG-AH 1
Radius Bd			1		1
Tibia Bd	1	2				2
Calcaneus GL		1

For the determination of the MNI of the juvenile reindeer in Kammern-Grubgraben, the tooth wear stage of Pd4, M1, and M2 were categorised by jaw and body side. The results for the season and the respective month for the Pd4 are summarised in Table 4.

Table 4

Kammern-Grubgraben. NISP of teeth present in the archaeological horizons of Kammern-Grubgraben, broken down by body side, jaw, tooth type, and tooth wear stage. The largest number of individuals detected (MNI) in each month is highlighted by grey shaded cells. Abbreviations, d: dextral, s: sinistral, Max: Maxilla, Mand: Mandibula
wear stage	Pd4 Max d	Pd4 Max s	Pd4 Mand d	Pd4 Mand s	wear stage	M1 Max d	M1 Max s	M1 Mand d	M1 Mand s	wear stage	M2 Max d	M2 Max s	M2 Mand d	M2 Mand s	M2 Max s
							KG-AH 1
4	1			3	2		1		2
5				2	3			1	2
6–8					4–5					1/2	1	1	1	3
							KG-AH 2
3	3	1	3	2	1
4	48	52	32	33	2	22	17	25	17
5	24	24	11	4	3	11	16	24	35
6–8					4–5					1/2	49	36	55	38	56

The calculation of the MNI of the reindeer at Kammern-Grubgraben yielded the following results: KG-AH 1 provided the remains of at least 11 reindeer (Table 5). No animals under six months of life were present, but five calves (juveniles) which died in the first winter of their lives, and two yearlings (subadults) from the second winter of life. The remains of at least four adults are present, two of which are females and one a male. In KG-AH 2, there are at least 165 reindeer individuals, including 90 calves from the first year of life and 49 yearlings, 11 adult females, 4 adult males and 11 indeterminable adults.

Table 5

Summary of the MNI calculated for reindeer in Kammern-Grubgraben
	KG-AH1		KG-AH2
age/sex	MNI	%	MNI	%
juveniles/calves	5	45	90	55
subadults/yearlings	2	18	49	30
adult males	1	9	4	3
adult females	2	18	11	7
adult ?	1	9	11	7
MNI total	11	100	165	100

[header 2] Comparison of species MNI between late MIS 3 to the early MIS 2

In the assemblages from the older phase (Fig. 5a, Table 6), the high MNI of 21 mammoths – the largest number of individuals in the data set – is particularly remarkable. At Kammern-Grubgraben, in contrast, only one mammoth is attested per layer, while reindeer is clearly dominant with an MNI of 165. This notable shift may partly reflect the abundance of these species in the landscape but seems to also signal a change in hunting preferences.

Table 6

Comparison of species and genera MNI between the (late) MIS 3 and (early) MIS 2 sites identified in the sites of the Early Gravettian and Early Epigravettian (Krems-Wachtberg: ¹Händel, 2017, 151; ²present study). * in the earlier studies no distinction was made between *Vulpes vulpes* and *Alopex lagopus*.
	MIS 3							MIS 2
species/genera	WA-AH 4.4¹	WA-AH 4.11¹	WA 1930¹	HU-AH 3.2¹	HU-AH 3.3/3.4¹	Langenlois A²	Pre-LGM total	KG-AH 2²	KG-AH 1²	LGM total
Bison sp.								2		2
Bos primigenius									1	1
Capra ibex	1	10	2	1		3	17	3	1	4
Cervus elaphus		1	1	3	1	1	7	2		2
Equus sp.	2	2		2	2		8	22	10	32
Colodonta antiquitatis	1	1		2	1		5
Ovibos moschatus			1				1
Mammuthus primigenius	2	2	8	7	2		21	1	1	2
Rangifer tarandus	2	2	2		3	3	12	165	12	177
Ursus arctos	1	2					3	1		1
Canis lupus	1	5	6	3			15	2	1	3
Gulo gulo	1	1	3				5	1		1
Alopex lagopus	3	6	5	1	1		16	1		1
Vulpes vulpes	*	*	*	*	*	1	1	1		1
Mustela sp.	1	1		1	1		4
Anser sp.						1	1	1		1
Branta bernicla					1		1
Lagopus lagops		1			3	1	5
Tetrao urogallus	1				2		3
Lepus sp.	2	3		2	2	1	10	5	2	7
Spermophilus sp.	1	2		1	1		5
Arvicola sp.	1	1		2			4
Citellus sp.								1		1
Cricetus sp.	1	1		1			3
Microtus sp.	1	1		6			8
Castor fiber				1			1
total	22	42	28	33	20	11	156	208	28	236

[header 2] Seasonal dating of the archaeological sites

Animal remains that can be used to determine the season were available for reindeer, horse, and ibex.

[header 3] Reindeer antler. For Kammern-Grubgraben, a total of five skulls of reindeer with attached antlers were identified. All other antlers of male animals used for artefact production were, as far as can be discerned, from shed beams (Pfeifer et al., accepted). Three un-shed antlers from KG-AH 2 were from female or young male reindeer. Processing marks on these antlers were not evident. Adult females and young males until 2 years of age shed their antlers in May (Herre, 1986, 205), indicating that the animals died during winter.

In addition, two skulls with fully formed attached antler fragments of male reindeer in KG-AH 2, showed partial traces of processing. Antlers of male reindeer are fully formed around the rutting season in autumn and shed thereafter (Herre, 1986, 205). These specimens thus either originate from animals hunted in late autumn or have been collected from naturally deceased reindeer. In any case, the observations on antlers do not allow any other seasonal dating than late autumn/early winter until the end of May at the most.

[header 3] Reindeer teeth. The analysis of thin sections of isolated teeth yielded no results, as the outermost layer was hardly or no longer preserved in most cases. Therefore, some teeth that were still in the bony alveolar ridge were embedded in resin. As a result, the diagnostic area was better preserved. However, this also only allowed a statement to be made about the animal's season of death on one tooth of an adult reindeer (maxilla M1, wear stage 6/g, according to Van den Berg et al., 2021, absolute scheme/Fig. 4). Although some imperfections can be recognised (remnants of the embedding resin, pieces of fragmented cementum), individual lines are visible (Fig. 6). The arrows point to the outermost line, which represents the last formation in the life of the animal. This is clearly recognisably dark, so that this specimen can be shown to have died in winter.

A further method used for determining the season was the combination of tooth dimensions and wear stage in the Pd4, as described above. In both jaws, there are no teeth with a wear stage lower than 3 (Fig. 7). In the teeth of both jaws, there is a clear tendency towards a decrease in crown height in connection with the wear stage. In the mandibular Pd4, there are hardly any differences in the dimensions between tooth wear stages 4 and 5, whereas there are clear differences in the maxillary Pd4 in these stages. In both jaws, a gap between tooth wear stages 5 and 6 is clearly recognisable.

As already observed in the Greenlandic caribou, Fig. 8 shows the temporal dimension in the connection between tooth wear stage and the tooth crown dimensions of the Pd4. However, probably not every tooth condition can always be defined precisely with a wear stage. In addition to a certain individual variation of the animals, also the presence of both sexes is expressed in their deviating dimensions. Despite these influences and the incomplete separation of individual groups, a development from top right to bottom left is recognisable, which represents the progress of time. The mean values ("M"), formed from individual tooth age stages (Table 7) also highlight this trend, where increasing tooth wear influences the crown height, but also the width of the tooth. Both dimensions decrease with increasing erosion and thus with increasing age. It is striking that despite the variation in the dataset, there is no overlap in the dimensions between tooth wear levels 3–5 and 6–8. This suggests that certain age groups are not represented in the sample. Presumably, this reflects the absence of animals that died during the summer months. Animals with a tooth condition of Pd4 "before eruption" and with tooth wear levels 0–2 are not represented. These tooth states would cover the period from May to around September. Animals with tooth wear stage 3–5 represent the first winter of life, animals with tooth wear stage 6–8 the second winter of life. Both, the absence of wear stages less than 3 and the gap or lack of overlap between 5 and 6 indicate that there was no hunting of young reindeer in the summer months. Tooth wear level 7 shows a wide variation, but two groups seem to emerge here, possibly representing male and female animals.

Table 7

Reindeer, Pd4 of the mandible and maxilla. Measurements (in mm) from both AHs (KG-AH 1 and 2) were used for the analysis. Mean values according to degree of tooth decay and presumed season, data see Supplement 4 and 5.
tooth wear stages	season	mandibula			maxilla
		heigth	length	NISP	height	length	NISP
3	late autumn (October)	9.73	24.18	4	10.05	19.10	2
4	early 1st winter (November)	9.10	23.78	31	9.03	18.19	33
5	high-late 1st winter (December-March)	8.51	23.84	29	7.29	17.73	17
6–8	2nd winter	5.17	22.38	32	4.18	16.32	13

Since tooth wear stage 5 in the Greenlandic caribou represents the time from around November onwards, the data for the reindeer from Kammern-Grubgraben can be further specified using wear stages 3–5. The animals with wear stage 3–4 therefore likely died before December. The Pd4 breaks through shortly after birth and according to Barboza & Parker (2008, 835) and Parker et al. (1990), reindeer neonates feed exclusively on milk for the first 4 weeks, as their stomachs are not yet fully developed at birth. For this reason, no tooth wear forms in the first weeks of life. From a certain age, calves begin to feed on plants in addition to milk. Data on weaning vary widely (Herre, 1986, 212). According to Herre (1986), this can occur as early as 2 months of age, but sucking has also been observed in 4–5 months old and even older calves. This means that tooth wear does not begin until late summer/autumn the earliest, when calves start to eat hard food alongside their mother's milk. Since tooth wear begins at stage 3 in Kammern-Grubgraben, a certain period must have passed by then after the start of feeding on hard food. The start of wear stage 3 cannot be precisely determined based on the available information, but it can be assumed that the first animals in Kammern-Grubgraben died around October. Since a certain difference between wear stage 3 and 4 is recognisable (Fig. 7), the animals of wear stage 4 probably died in November. Consequently, the animals with wear stage 5 died in mid-winter, and in comparison to the data of the Greenland caribou, around December to March. As explained above, the gap in the data indicates that the animals with wear stages 6–8 died in the second winter of their lives.

To summarise, it can be stated that reindeer were hunted exclusively in winter in Kammern-Grubgraben in all documented periods.

[header 3] Reindeer fetal bone. One reindeer fetal bone was identified from KG-AH 2. This radius had a shaft length of 39.24 mm (Fig. 9a). The determination of the season based on measurements in reindeer fetals cannot be given due to the lack of studies and data. Spiess (1979) gives calculations for the hind foot, or for the individual bones in the hind foot and for the humerus, but not for the radius. Therefore, the season of the fetal stage of the bone from Kammern-Grubgraben is narrowed down with the help of comparative skeletons (Fig. 9b). The comparative radius from a parturient caribou was 105.10 mm in length and almost three times as long as the Kammern-Grubgraben find. In this comparative skeleton, the crown tip of the Pd4 was broken through the bony crest. A breakthrough takes place shortly after birth (see Table 1), but it is likely that there is some variation in the breakthrough time, because the crown tips of this skeleton had already appeared. Thus, the comparative animal may have died around the end of May. Due to the size of the radius, the fetus from Kammern-Grubgraben was many weeks younger than the modern comparative skeleton. A precise indication of a certain month cannot be given, but the season for this fetal bone can at least be limited to a period from mid-winter to the beginning of late winter.

[header 3] Horse teeth. The analysis of tooth thin sections of horses yielded no result, but several milk incisors gave indications about the season. Two left maxillary Id3 from KG-AH 2 are certainly from two different animals. Their occlusal surfaces were very slightly worn (Fig. 9c-d). If they have been born between end of April and begin of July (Volf, 1996, 94), the animals died in winter of their second year of life, which was in months November to February at the latest.

[header 3] Ibex jaw. The vast majority of ibex remains came from adult animals that did not provide any information about the season. The analysis of tooth thin sections yielded no result. Only a few skeletal elements came from young animals, including a mandible with milk teeth from KG-AH 2 (Fig. 9e). The milk teeth were slightly to moderately chewed off. Although the M1 was no longer present, the tooth alveolus showed that the tooth had already erupted. According to Nivergelt and Zingg (1986, 399), the permanent molars appear at 15–30 months. Since the birth of ibexes - based on data from modern populations (Nivergelt & Zingg, 1986, 399) - usually takes place in the first three weeks of June, this animal probably died in the second winter of its life. The data on the season at Kammern-Grubgraben therefore shows that not only reindeer in both AHs (KG-AH 1 and 2), but also ibex and horses (verified in KG-AH 2) were hunted exclusively in winter.

[header 2] Comparison of the seasonal occupation between the sites of the pre-LGM sites and Kammern-Grubgraben

Several fauna elements from Krems-Hundssteig can be used for seasonal dating for the various occupation events and revealed settlements at all times of the year (Fig. 10). The postcranial element of a foetal or neonatal reindeer (Händel et al., 2021b) is the only seasonal indicator at Krems-Wachtberg and is originally determined as a winter settlement (Fladerer et al. 2014). But the season is more likely a spring occupation, as a bone in the size of a ready-to-birth reindeer indicates a dating around the birthing season (May/June). The other layer from this site considered here is WA-AH 4.11 which is again a palimpsest.

At Langenlois A, a winter to spring occupation (November to mid-March) is attested based on a single find only, the skull antler of a female reindeer. However, it does not allow a more precise limitation of the season. This site is younger than the Krems sites.

For Kammern-Grubgraben, the season can be determined based on ibex and horse teeth, and on reindeer antlers, fetal bones, postcranial elements, and teeth, which all clearly indicate human presence exclusively in winter. The main season of occupation in both archaeological horizons, KG-AH 2 and KG-AH 1 can be defined as winter. The evidence of settlement extends over the months October to March based on the Pd4. It is possible that the marginal months of the representation (October and February/March) are based on a certain variation in the basic data of the reindeer individuals, and that the main settlements took place around November-January. However, it is also possible that the settlement always or just temporarily extended over these months.

In summary, the data so far indicate that the seasonal settlement pattern in the region appears to have changed from occupations during different seasons in the Gravettian to a winter settlement only in the LGM.

[header 2] Reindeer prey pattern at Kammern-Grubgraben

The age and sex composition of the reindeer assemblage from Kammern-Grubgraben is remarkable. The largest proportion with more than half of the finds are calves of around six months of age, the second largest with about one third are yearlings from the second winter of their lives. Among adult reindeer, there are about three times as many females as males (Table 5).

This share of calves, yearlings, and adults in the find spectrum of Kammern-Grubgraben are not typical for the shares in a herd of reindeer (e.g., Herre, 1986, 211; Heptner et al., 1966), albeit Davis et al. (1980, 597) mention that it is not easy to determine the sex and age structure of reindeer herds. In general, different proportions can be observed in the individual groups of all herds, which vary depending on the season or even between different years. The influence of diseases, predators and hunting by humans affects the composition in different ways.

Sexually mature female reindeer usually give birth to one calf per year (Bergerud, 1980, 556; Herre, 1986, 210). This means that the proportion of adult females to calves should be the same. Some of the adult animals (7%) could not be distinguished by sex (Table 3). If these were considered, the proportion of adults would increase. If all indeterminable animals were assumed to be female, their proportion would increase to 14%, but still be significantly underrepresented compared to the calves, with more than 50%. However, calf mortality is high in the first few months of life, so that by the first winter, a part of the calves has died (Bergerud, 1971; Bergerud et al., 2008; Mahoney et al., 2016; Wittmer et al., 2005). In North American caribou herds, the mortality rate of calves is frequently observed to be as high as 80–90% (Mahoney et al., 2016, 96). Most of these deaths occur within the first few months of life, with predation identified as the leading cause of mortality (Bergerud, 1980, 556). Large herbivores often experience significant variation in juvenile survival rates (Gaillard et al., 1998). In the case of caribou, predation on juveniles and low recruitment are consistently acknowledged as significant factors contributing to population declines (Boertje et al., 1996; Wittmer et al., 2005; Latham et al., 2013). Wildlife biology studies on Newfoundland caribou showed that 50% of the calves died within the first six weeks during a population decline (Mahoney et al., 2016). In contrast, 50% of the calves only died in the first seven months of life during a population peak. In a study by Mahoney et al. (2016), the main cause of calf mortality was medium to large carnivores (coyotes, lynx, black bears). According to reports, foxes and ravens are significant contributors to the mortality of newborn caribou in Canada (Nowosad, 1975, 207). The involvement of ravens and sea eagles in the deaths of calves has been confirmed in Siberia (Baskin & Danell, 2003, 149). Thing and Clausen (1980, 434) documented a mortality rate of 50% for 2 to 3-month-old calves in the Kangerlussuaq area in Greenland in 1977. This elevated percentage was attributed to various illnesses affecting calves, including infections, parasitic infestations, and diarrhoea. These health issues were often linked to the compromised physical condition of pregnant mothers, resulting in the weakening of the fetal organism and increased vulnerability of the young animals to these ailments (e.g., Espmark, 1980, 495). Miller and Broughton's (1974) research in Canada indicated that a significant proportion of calf deaths resulted from severe weather conditions or instances of desertion or separation from their mothers (refer to Skogland, 1989, 55). McEwan's findings in 1958 (McEwan, 1959) reported a calf mortality rate of 33.5% among Canadian Barren-ground caribou from birth to August. Most of these deaths occurred shortly after birth on the calving grounds and were attributed to adverse weather conditions (Kelsall, 1968, 165).

Even if a peak in the reindeer population and favourable conditions are assumed for the time of settlement in Kammern-Grubgraben, there should be significantly fewer calves in the population than have been identified. If one calf per adult female and a certain loss of young animals during winter is assumed, calves are particularly overrepresented in Kammern-Grubgraben compared to adult females.

A higher mortality rate affects male rather than female reindeer from the juvenile stage at the latest, which progresses with increasing age. Semenov-Tian-Shankii (1975, 159–160) found out, that in kills by bears, males predominate over females (10:3) and older individuals over younger (6:1). Males and old reindeer, being less wary than females and younger animals, are most likely to be killed. As a result, the number of male reindeer decreases with increasing average age in a herd (e.g., Heptner et al., 1966 in Herre, 1986, 211; Pasda, 2009, 107–108). In Canada, for example, the relationship was 63 bulls : 100 cows in 1968 and in 1970, and 45 bulls : 100 cows between 1975–1978 (Davis et al., 1980, 595). A predominance of females, as can be observed in the case of the adult reindeer from Kammern-Grubgraben, is therefore to be expected.

To further illustrate that the proportions of young animals in Kammern-Grubgraben in no way correspond to a natural composition of a reindeer herd, two examples of recent, undomesticated herds are given. The first is from Canada: The Nelchinas herd was documented twice at intervals of 10 years (Bos, 1975; 1960 Fig. 11b, and 1970 Fig. 11c). Although the proportions of each sex and age group vary somewhat between the two different years of the study, a similar herd composition can be seen: a particularly large number of adult females lived in the herd, few males, and many juveniles. In both years, the adult females made up almost half of the herd.

The second example is a Siberian reindeer herd that was documented (Heptner et al., 1966 in Herre, 1986, 211) in autumn (Fig. 11d) and in April (Fig. 11e). This comparison clearly shows the mortality of young animals in their first year of life. If we assume that every adult female reindeer had a calf in spring, then around 25% of the calves in this herd had already died by autumn. The mortality trend in the proportion of juveniles continues until April. Due to the imprecise data of the yearlings in the subadult stage (Heptner et al., 1966), it should only be noted here that the overall number also decreases at this age stage. In the adult stage, the proportions of males and females differ only slightly between autumn and April. However, the ratio of males to females is significantly different at both times and varies between a ratio of around 1:2 (Fig. 11b) and 1:5 (Fig. 11c).

A direct comparison of the data on the reindeer represented in Kammern-Grubgraben (Fig. 11a) with the data on the population structure of the Canadian (Fig. 11b-c) and the Siberian herd (Fig. 11d-e), shows that the proportion of calves is significantly higher in Kammern-Grubgraben. The proportion of subadults (yearlings) is slightly higher and the proportion of adults is significantly lower.

The data from the reindeer finds from Kammern-Grubgraben therefore show clearly different representations of individual groups in comparison with modern reindeer herds with a striking high number of calves, followed by a higher number of yearlings and by the low number of adult animals. The ratio between female and male adult animals of around 1:3, on the other hand, corresponds to a natural ratio.

[header 2] Significance of reindeer hides and fur based on ethnographic sources

The results on the significance of reindeer in Kammern-Grubgraben show a substantial change in people's behaviour. The representatives of the genus Rangifer, i.e., reindeer and caribou (in the following, only referred to as reindeer) have served as a significant resource for numerous human communities across northern Eurasia and North America for millennia (Burch, 1972). Prehistorians often depict the species as offering a plentiful, readily accessible, and consistently dependable resource for humans. While a sufficient supply with meat and fat is usually considered essential for humans in an arctic environment and for Palaeolithic hunter-gatherers, the importance of reindeer skins is comparatively underestimated. However, the fur is one of the most important parts of a reindeer body for people living in an arctic environment (Pasda, 2013a, 2013b; Pasda & Odgaard, 2011). In the historic past, it is known that obtaining fur was sometimes the more important reason for hunting than the reindeer’s meat or fat (Pasda, 2022, 269–270).

From the age of approx. 1.5 years, reindeer have a remarkable ability for storing body fat during times of abundance (summer), to be utilised later in times of scarcity (winter) (Nilssen et al., 1984). In fact, more than 30% of carcass weight is composed of fat in autumn, and it has been suggested that fat deposition is a factor of crucial importance for survival of this species (Reimers & Ringberg, 1983). This ability to store fat may have been particularly important for humans during the cold season and therefore also a reason for the intensive hunting of reindeer in winter. Due to the clear sexual dimorphism, adult male reindeer have a larger body mass and thus provide more meat and fat than females. However, the proportion of female to male animals in Kammern-Grubgraben corresponds to a natural ratio, so that these results do not indicate selective hunting for the primary purpose of obtaining a lot of food. However, the osteoarchaeological results show that especially calves from the first winter of their life and besides yearlings probably from the second winter are represented at Kammern-Grubgraben. This fact, in conjunction with the information on the significance of reindeer fur for arctic people, leads to the conclusion that a major purpose of the camp at Kammern-Grubgraben was obtaining and processing fur. The high degree of bone fragmentation in Kammern-Grubgraben shows that in addition to their hides the entire animals were exploited for meat, fat, and other products. In this context, the massive stone structures documented at the site and interpreted as meat caches (Einwögerer, 2021), might have served as storage facilities to protect a surplus of meat that could not be consumed directly. The results of our investigations therefore provide clear indications of intensified reindeer fur use, presumably as an indication of adaptation to a cold environment.

Reindeer fur has special properties. Reindeer are perfectly adapted to survive in an arctic environment. They can cope with a small amount of food and can survive in very cold temperatures. To achieve this, reindeer and caribou have developed some unique physical characteristics.

First, their fur is very dense. Reindeer have over 5,000 hairs per cm². In comparison, humans have about 600 hairs per cm². Furthermore, reindeer hairs are hollow (Timisjärvi et al., 1984, 601). Their interior is filled with air, a characteristic that allows them to perfectly isolate the body from the icy temperatures which can reach − 50°C. The animal possesses a gentle, fine underlayer of woolly insulating hairs, complemented by a longer, coarser outer coat of protective guard hairs (Cuyler & Øritsland, 1999, 93). The longer guard hairs help to shed rain and the worst weather. The winter coat is particularly warm. Winter insulation of Svalbard reindeer was approximately three times that of summer. Studies on wind protection prove particularly good protective properties, especially for winter skins (Cuyler & Øritsland, 1999, 93; Hammel, 1955). These characteristics causes the fur to be especially warm, wind protecting and waterproof.

Ethnographic studies show the great importance of reindeer skins as insulation against the cold and protection against wind and moisture for all communities in arctic regions (Banfield, 1951; Crow, 1974; Hammel, 1955; Burch Jr., 1972; Moote, 1955). Between the individual modern polar tribes, the use of reindeer skin as well as the structure of the clothing is remarkably similar everywhere (Latreille & Orsenna, 2019, 201). Hides and fur are used as tent roof, as warming sledge blankets and as a floor underlay in the winter tents (Graves and Gunn, 1991, 95), as an outdoor seat pad (Latreille & Orsenna, 2019, 57), as sleeping bag (Latreille & Orsenna, 2019, 37), as coat or child carrier, or for gloves or caps (Latreille & Orsenna, 2019). The back extension of women's coats served as a carrier for their babies and was lined with a reusable "nappy" made of reindeer skin or moss (Alia, 1991, 101–102). The fur is particularly important for shoes, not only in cold, but also in wet conditions as the hair is waterproof. Due to their property of being waterproof, they are for example preferred for boots in the Finnsko style (Norv.: Finnsko, Engl.: Finnesko) developed by the Sami people which are made of fur from the legs of reindeer (Riffenburgh, 2006, 168–169; Alia, 1991, 101–102).

There are strong seasonal differences in the condition of the fur, so that skins from different seasons and from different animals are used for different purposes. The change of hair in reindeer begins in March with the shedding of the undercoat, which reappears in May (Herre, 1986, 205). This is followed by the outer hair („awn hair“), which is most intense in June. The coat is completely renewed in July and August, but then increases in length until winter.

Reindeer skins have different characteristics depending on age, sex, and season (Oakes & Riewe, 1996, 40). According to the hunters, leg skins from caribou killed in spring are rarely used because the intermediate claw bag (Sinus interdigitalis) enlarges at this time, which reduces the quality and size of the usable skin parts (Webster, 1949 in Oakes & Riewe, 1996, 40). The skins of caribou hunted from mid-July to mid-September are used for clothing, short-haired skins from July for stockings and August skins with longer hair for boots, so-called Kamiks (Webster, 1949 in Oakes & Riewe, 1996, 40). In Alaska, the skins of old bulls killed around October were used for the soles of overcoat slippers and Kamiks, as they have thicker skin. At first, however, the skins of bulls are used as sledge covers for several months. The ropes on the moving sledge rub against them and make them soft (Oakes & Riewe, 1996). Leg skins from caribou shot from October to November are favoured for adult boots, and the leg skins from calves are preferred for children's boots. In terms of thermal insulation, there is no big difference between young and older reindeer. At the same time, there is no significant difference between calf and adult insulation for the summer, autumn, and winter seasons (Cuyler & Øritsland, 1999, 93). Calf fur insulates as well as adult fur. According to modern fur specialists, the blankets of four- to six-month-old calves, however, are preferred for fur processing in modern times (Dathe & Schöps, 1986, 279–281). But even calves that are up to one year old still provide fine, sometimes silky skins. In 1965, the amount of fur required to make a so-called "body" coat, a Pijiki coat, was given as 16 to 22 young animal skins (Schöps, 1965, 7–12). This was based on a sheet with a length of 112 centimetres and an average width of 150 centimetres and an additional sleeve section. This corresponds roughly to a coat material for a slightly flared coat in size 46 for 2014. The maximum and minimum fur numbers can vary due to the different sizes of the sexes of the animals, the age groups, and their origin. Depending on the type of fur, these three factors have different effects.

Although other sources emphasise the use of winter skins or more precisely winter leather (Oakes & Riewe, 1996), Stenton (1991) suggests that in late autumn and throughout winter, the hair of adult animals is too thick and the skins too heavy for use as clothing. Instead, the pelts from winter kills are used for sleeping coats/blankets and other purposes. The best blankets (coats) of adult animals are those from autumn and the beginning of winter, as autumn coat is shorter and less brittle (Damm, 1950, 19). Skins of young reindeer are categorised as "fine" within the modern hair fineness classes (silky, fine, medium-fine, coarser, hard; Schöps & Häse, 1955, 39–40). According to fur experts (Dathe & Schöps, 1986, 279–281), the blankets of four to six-month-old calves are favoured for fur processing in modern fur processing facilities. That would mean that these skins come from calves from autumn or winter. As mentioned, there is no difference between young and older reindeer when it comes to heat insulation (Cuyler & Øritsland, 1999), however, young reindeer skins are particularly silky in winter and concerning to ethnographic sources this skin is preferred for clothes of the children (Alia, 1991, 101–102).

As for the consumption of fur by polar communities living as nomads, there are different reports about the number of reindeer skins needed per year. One Russian source gives the number of 300 reindeer for a family of 6 people for North Asian nomads, to be able to live from it "with some comfort and in moderate prosperity" (Schmidt, 1970, 378–380). However, the fur has a poor durability caused by the brittle grain hairs. The durability is given as "one year at most" (Dathe & Schöps, 1986, 279–281; Stenton, 1991). That means that there is a constant need to renew the clothes and thus to obtain new hides.

The information on reindeer skins can be summarised as follows: Reindeer skins are particularly warm, wind-insulating, and water-repellent. According to ethnographic sources, they are and were used intensively in arctic regions for various purposes such as clothing and household items. Depending on the season, age and sex of the animals, the furs serve different purposes. Winter pelts are particularly warm, however, the winter coats of adult animals are too thick to produce clothing, while the pelts of calves and young animals are very warm and in addition especially soft and silky. However, as the durability of the fur is low, they must be constantly renewed.

Our results demonstrate that a shift of availability of huntable animals due to climatic changes has taken place in Lower Austria between the pre-LGM and the LGM. Changes are also evident in the local paleoecology (Reiss et al., 2023), in the season of occupation, and in the material culture (Pfeifer et al., accepted.). Our earlier studies proposed more favourable climatic conditions for Krems-Wachtberg as opposed to Kammern-Grubgraben (Maier et al., 2021; Reiss et al., 2023). Nevertheless, high-resolution δ¹⁸O analyses of horse teeth do not reveal substantial climatic differences between the pre-LGM and LGM as during both periods a polar tundra climate with largely similar cold temperatures during stadial conditions prevailed (Reiss et al., 2024). However, there was a prolonged cooling phase, the GS 3, in which environmental and faunal changes must have occurred in the study area. This is evidenced by the strong reduction of ecological herbivore niches from the pre-LGM to the LGM (Reiss et al., 2023). A large-scale comparison between populations west and east of the Alps indicate complementary ecological adaptation with preferences for non-permafrost environments in the west and permafrost environments in the east (Maier et al., 2016). Living in permafrost environments comes with special requirements for clothing, which must perform particularly well in terms of isolation and protection against the cold.

The manufacture of tailored clothing was certainly already important before the LGM, and various pointed implements of osseous materials from Krems-Wachtberg (Einwögerer et al., 2008) may relate to skin processing. However, it is during the peak of the LGM, at around 23 ka cal BP, that eyed bone needles appear simultaneously in the Epigravettian in Eastern Europe, the Badegoulian in Western Europe (Pétillon & Chauvière, 2017) and at Kammern-Grubgraben (Pfeifer et al., accepted). The wide dissemination of this innovation was probably the result of re-established supraregional contacts during that time window and is also attested in the lithic industries (Händel et al., 2021a; Terberger, 2013). Eyed needles are suitable to produce very regular and tight seams on clothes, gloves, and shoes (d’Errico et al., 2018), which was particularly desirable during the LGM (Terberger, 2013). Moreover, they were certainly also used to apply decorative objects on clothing, such as perforated animal teeth and perforated molluscs and/or molluscs with natural holes. In particular fossil dentalia are very abundant at Kammern-Grubgraben (Neugebauer-Maresch et al., 2016).

As emphasised previously (Stenton, 1991), winter clothing requirements was not only a necessary precondition for the successful operation of traditional Inuit economic systems during winter and spring but should be considered a vital component in modelling subsistence and settlement behaviour of prehistoric people in an arctic environment. Stenton (1991) points to the problem that despite the array of technological advancements developed by prehistoric communities to cope with arctic environmental challenges, it is surprising that clothing has received minimal attention from archaeologists. While excavations have yielded fragments and occasionally intact pieces of clothing (Maxwell, 1985; McCullough, 1989; Park, 1986; Newell, 1984; Hansen et al., 1991), the topic is typically addressed indirectly, often through references to specific types of sewing tools and skin preparation instruments (such as bone needles, awls, needle cases, thimbles, microblades, and skin scrapers), or evidence of faunal remains (Maxwell, 1985; McGhee, 1978). Both archaeological evidence and ethnographic parallels indicate that crafting fur garments suitable for arctic conditions was a labour-intensive process demanding considerable skill. Therefore, the creation of tailored fur clothing represents a pivotal adaptation of the human species to circumpolar regions and in Palaeolithic periods.

Acknowledgements

We gratefully acknowledge financial support from the German Research Foundation (DFG) and the Austrian Science Fund (FWF). The entire method of creating thin sections was explained and supervised by Dr. Carsten Witzel and Dr. Patricia Kahle, University of Hildesheim. Without their intensive support and friendly assistance, the successful creation of the thin sections would not have been possible! We also thank U. Simon for valuable discussions on the study sites.

Authors Contribution

K.P. taxonomically identified, recorded, and analysed the studied animal remains from Kammern-Grubgraben and Langenlois A, wrote the original draft and prepared all figures and tables. L.R. provided paleoclimatic background data. S.P. analysed the organic material of the sites and wrote the text part for the sewing needles. L.R. and W.S. produced the decalcified thin cuts and K.P. the resin dental thin sections, W.S. documented the samples. T.E. and M.H. provided the study material. K.P. drafted the study design with contributions of C.M., A.M., T.E., M.H., and N.B.. All authors contributed substantially to the text and reviewed and approved the manuscript.

Funding German Research Foundation (DFG) (grant AZ. MA 4235/12-1, PA 1616/8-1, 1616/8-2; project no. 424736737 & grant PF 841/4-1 project no. 442218485) and Austrian Science Fund (FWF) (project I-4306-G25).

Data Availability Data is provided within the supplementary information files.

Ethical Approval

Not applicable

Competing interests

The authors declare no competing interest.

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Ice Age apparel – changing prey patterns towards the Last Glacial Maximum and the role of reindeer fur for clothing at Kammern-Grubgraben

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Abstract

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Introduction

Methods

[header 2] Dental cementum analysis

[header 2] Odontochronology and tooth wear of reindeer

[header 2] Sex determination of adult reindeer

Results and Discussion

[header 2] MNI calculation for Kammern-Grubgraben

[header 2] Comparison of species MNI between late MIS 3 to the early MIS 2

[header 2] Significance of reindeer hides and fur based on ethnographic sources

Conclusion

Declarations

References

Additional Declarations

Supplementary Files

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