The fossil cave hyena of Goyet, Walsin and Hastière (Belgium): osteometry and taphonomy

Transcription

1 FACULTY OF SCIENCES Biology Department Palaeontology Department Academic year The fossil cave hyena of Goyet, Walsin and Hastière (Belgium): osteometry and taphonomy Debora Beke Thesis submitted to obtain the degree of Master in Biology Supervisor: Prof. Dr. J. Verniers Supervisor and tutor: Dr. M. Germonpré (RBINS)

2 May 2010 Faculty of Sciences Biology Department All rights reserved. No part of the publication may be reproduced in any form by print, photo print, microfilm, electronic or any other means without written permission from the publisher.

3 Table of contents 1. INTRODUCTION General introduction The spotted hyena - Crocuta crocuta Recent spotted hyena Fossil spotted hyena or cave hyena 2 2. AIMS 2 3. MATERIAL AND METHODS Description of the sites Description and organisation of the material Methods and measurements Cranial measurements Postcranial measurements Age determination Weathering and gnawing traces Database Comparing fossil and recent spotted hyena RESULTS Inventory of the fossil material Description of the fossil material Cranial Postcranial Age determination Trace fossils Coprolites Other traces Weathering 47

4 4.6. Inventory of the recent material Description of the recent material Cranial The fossil and recent spotted hyena DISCUSSION Taphonomy Age determination Use as communal dens? Gnawing traces and cannibalism Fossil and recent spotted hyena CONCLUSIONS DUTCH SUMMARY THANKS REFERENCES 76 ANNEXES 79 Alphabetical List of abbreviations 79 Statistical tests Primary data CD-Rom CD-Rom

5 1. INTRODUCTION 1.1 General introduction In Belgium many local deposits of Late Pleistocene fossil mammals are found in the Flemish Valley: a complex of valleys eroded and refilled by the activity of rivers estuaries and eolian action (GERMONPRÉ, 1995 and 1996). In the Ardennes, fossil mammals are very well represented in caves along the Meuse River (GERMONPRÉ, 1995, 1996). In this research we present the osteometric and taphonomic characteristics of bone assemblages from the fossil spotted hyena found in three caves: the Goyet cave, Trou de l hyène (Walsin) and the Hastière cave. These three caves situated in Namur province, were excavated by Dupont between 1860 and Dr. Germonpré, associated with the Palaeontology department of the Royal Belgian Institute for Natural Sciences (RBINS), has already done research on the Goyet cave. Studying hyena taphonomy can be of great importance for paleontological research as hyenas produce large osseous assemblages (POKINES & PETERHANS, 2007). 1.2 The spotted hyena Crocuta crocuta The recent spotted hyena Recent spotted hyenas are found in the sub-saharan region of Africa and are distinguished by their exceptionally enlarged premolars, robust skulls and teeth (BINDER & VALKENBURGH, 2000). They live in social groups (clans) of 5-90 members that recognize each other by sight, scent and vocalisation. Adult females, cubs and immigrant adult males live together (KRUUK, 1972; HOLEKAMP & SMALE, 1998). Adult females bear litters of one or two cubs, born in isolated natal dens with fully opened eyes and fully erupted canine and incisor teeth. After three to four weeks the cubs are brought to a communal den where all cubs live together until they are eight to nine months old. After this period, they leave the communal den and start to explore the world outside; sometime later weaning starts. Eventually they will reach adult and reproductive maturity (at 24 months for males, at 36 months for females) (KRUUK, 1972; HOLEKAMP & SMALE, 1998). Holekamp and Smale (1998) described this hyena life through five developmental stages; each starting and ending at a specific age and characterized by important moments in life. Cannibalism among spotted hyenas is a controversial issue. According to Kurtén (1968) spotted hyenas are not intentional cannibals as eyewitnesses state that hyenas do not eat other hyenas. Kruuk (1972) on the contrary states that protection against other hyenas is a necessary behaviour pattern and that hyenas kill and eat almost everything, including members of its own clan. He proposes that this is one of the reasons why females are bigger than males: to keep the latter away from the cubs. Also the ability of the cubs to creep out of reach of the adults may be a protection against cannibalism, as well as the fact that females always keep their litters in close proximity to each other so that cubs are protected when some mothers are away. 1

6 1.2.2 The fossil spotted hyena or cave hyena During the Pleistocene, the cave hyena was widespread in Eurasia and is therefore represented in many European bone caves. These caves were often used as dens (KURTÉN, 1986; KLEIN & SCOTT, 1989). Cave hyenas are distinguished from the now living African spotted hyenas mainly by size and limb proportions. They were carnivores with powerful bone cracking teeth with carnassials forming an effective shearing device; the canines were relatively feeble (KURTÉN, 1965, 1986). The cave hyena is described by many authors as a subspecies of the spotted hyena (Crocuta crocuta spelea). Rohland et al. (2005) did a DNA-analysis on Pleistocene and recent spotted hyenas and came to the conclusion that the sequences of both clades are intermixed, which suggest that both clades are not monophyletic. Furthermore, there were at least three migrations of African spotted hyenas to Europe and Asia around 3, 1 and 0 MYA. These last two migration events gave rise to the European populations (ROHLAND et al., 2005). At the end of the Ice Age, the cave hyena became extinct in Eurasia. As the cave hyena and recent spotted hyena are closely related, we can use our knowledge of the recent spotted hyenas to make interpretations about the way of life of cave hyenas. In general, we presume that their life was characterized by similar life strategies, that is, cubs are also raised in communal dens. 2. AIMS The fossil hyena material from the three different Belgian sites will be studied and measured in order to answer some questions. What kind of skeletal elements are found and which ones are lacking? Are there any marks present on the bones, like gnawing traces that could point to cannibalism? Do we find significant differences between the caves in the foregoing respects and why do we find differences? The main question is whether the hyenas used the caves as communal dens or whether they used them just as safe places to shelter. Do we find, besides adult bones, juvenile remains that can indicate the use of a cave as communal den? We also compare the fossil material of Crocuta crocuta with some recent material. This comparison may reveal significant differences between both groups. 2

7 3. MATERIAL AND METHODS 3.1 Description of the sites Goyet, Walsin and Hastière are situated in the Namur province in Belgium (fig. 3.1). Fig. 3.1: Location of Goyet, Walsin and Hastière. (2010 Google Data 2010 Tele Atlas Europe Technologies, PPWK) GOYET Goyet is situated at the confluence of the rivers Samson and Strud. On the right bank of the Samson lies a limestone cliff which harbours a series of caves. Those caves were excavated by E. Dupont between 1868 and Dupont numbered the different caves, five in total. The hyena material from Goyet analysed in this study was found in the third cave. The location of this third cave is situated 15 m above the Samson, with its entrance to the southwest. The cave is very long and connected with the other caves trough a series of galleries (GERMONPRÉ, 1996, 2001). Dupont divided the cave in three parts: Chamber A, B and C (fig 3.1). Chamber A is about 25m deep and 5m wide; chamber B is connected with A through a small gallery and is about 10m in depth. Chamber C lies the furthest from the entrance ( m) (GERMONPRÉ, 1996, 2001, 2004). 3

8 Dupont described five successive bone horizons in this third cave, each horizon separated by sterile alluvial sediments. As mentioned in the introduction, Dr. Germonpré (1996, 2001, 2004) has already started to study the Pleistocene assemblages: Bones from horizon 1 were mostly found in chamber A. This horizon yielded remains from herbivores and carnivores, but also a few human bones (Germonpré, 1997). In this horizon 1261 identified bones and 700 unidentified remains were counted (Germonpré, 2001). On several bones from this horizon AMS dating was done; the uncalibrated ages are listed in table 5.1. The bones from the muskox and the horse were modified by prehistoric man (Germonpré, 1996, 1997, 2001). Species Calibrated Age (yr BP) muskox /- 90 horse /- 90 cave hyena / /- 400 cave bear Table 3.1: Uncalibrated ages from muskox, horse, cave hyena and cave bear; Goyet- horizon 1 (Germonpré, 1996, 2001). The cave bear and cave hyena bones were located deeper in the chamber and have another origin than the human modified bones of the muskox and the horse that were found at the entrance of the chamber. The different dates of the hyena bones point to the mixed nature of the horizon (Germonpré, 1996, 2001, 2004). Horizon 2 of Goyet is also found in chamber A; a total of 1706 identified bones and several hundred unnumbered remains were collected. It was separated from the first horizon by sterile deposits with a thickness of 10 to 15 cm (GERMONPRÉ, 2001). Horizon 3 of Goyet is the richest horizon from chamber A: some 3700 identified bones and hundred unidentified ones were excavated. This horizon was separated from the second one by sterile deposits with a thickness of 10 to 30 cm (GERMONPRÉ, 2001). Horizon 4 is found in the back of chamber A, in chamber B and is the only bone horizon present in Chamber C but Dupont mainly describes this horizon to chamber B (GEMONPRÉ, 1997, 2004). The majority of the bones from horizon 4 come from cave bears, followed by remains from horse, reindeer, hyena, bison, wolf and some other species. Some human remains from an adult and a child were also found (DUPONT, 1906; GERMONPRÉ, 2004). 4

9 Fig. 3.1: Map of chamber A, B and C from the third cave of Goyet (Germonpré, 2004) WALSIN Walsin is situated near the river the Lesse. Several caves were found along its valley of which two situated in Walsin: Trou de l ours and Trou de l Hyène. We used the fossil hyena bones found in Trou de l Hyène which will be referred to in what follows as the Walsin cave. The cave was located 10m above the right bank of the river but was destroyed in 1893 by the construction of a railway. It was a low and narrow gallery with a double exit, covered with a deposit of sandy clay rich in bones. Under this bone layer, a layer of stalagmites was present and this layer in turn was situated above yellow clay blocks in which bones of reindeer, horse and two flints were found. The bone layer yielded 16 different species, including hyenas and was excavated by Dupont in 1866 (DUPONT, 1905). HASTIÈRE The Hastière cave lies in the ravine Tale des Tahaux, a tributary of the river Meuse in Namur province. A wide cave entrance leads to the centre that was filled with stones and loam. Five horizons could be distinguished in which a lot of fossil mammals and some human remains were found. The human remains were limited to horizons 1, 2 and 3; the several faunal remains were listed by Dupont in 1872 during the excavation (DUPONT, 1906; EHRENBERG, 1935). According to Dupont (1906) these remains indicate that this cave had a succession of inhabitants: it was a lion den, a hyena den, a cave bear den and also occupied by prehistoric men. Unfortunately little published information on this cave is available. Besides some unpublished notes made by Dupont in , the faunal list of Dupont, published by Ehrenberg in 1935, is the only available information about the excavation. For this study we use the fossil hyena material of horizons 1, 2 and 5. 5

10 3.2 Description and organisation of the material Dupont identified the fossil material and marked every bone with a little tag upon which he mentioned the species, the skeletal element and its position (left or right). Then he sorted all the skeletal elements per species and per type on plaster plateaus. Each plateau received a number and a small card with a brief description, a date and Dupont s initials. The entire collection is kept at the Royal Belgian Institute of Natural Sciences (RBINS) in Brussels at the Palaeontology Department. Except for one skull, all the fossil material belongs to this Dupont collection and was analysed and measured at the institute. The fossil skull is kept at the Research Unit Palaeontology of the University of Gent. It was found in the North Sea. Besides fossil material, also some recent material is used for a comparative study (see section 4.6). This bone material is also kept at the RBINS, Vertebrates Department. Every piece has a tag with information about the find: year, place, sex, etc. 3.3 Methods and measurements First, we checked if the determination of Dupont was correct. Sometimes a skeletal element received a wrong description. In order to check the determination, we used atlases by Reynolds ( ), by Pales & Lambert (1971) and Pales & Garcia (1981). Then we gave each element a number added to the plateau number. Sometimes elements were already numbered and it happened that different elements of one plateau had the same number. In such case we added a letter. Dupont did not leave any osteometric data and our next task consisted of measuring most skeletal elements. Measurements were made according to von den Driesch (1976) with a slide ruler with an accuracy of 0.01 mm. The skull from the North Sea could not be measured with the same slide ruler and was measured with an accuracy of only 1 mm Cranial measurements The numbers refer to the numbers of the skull and jaw measurements proposed by von den Driesch (1976). A. SKULL-MEASUREMENTS (1) total length: from Akrokranion to Prosthion (2) condylobasal length: aboral border of the occipital condyles to Prosthion (8) viscerocranium length: from Nasion to Prosthion (12) snout length: from the oral border of the orbits (median) to the Prosthion (15) length of cheektoothrow 6

11 (25) greatest breadth of the occipital condyles (27) greatest breadth of the foramen magnum (28) height of foramen magnum (from the Basion to the Opisthion) (29) greatest neurocranium breadth = greatest breadth of the braincase (34) greatest palatal breadth: measured across the outer borders of the alveoli (35) least palatal breadth: measured behind the canines (36) breadth at the canine alveoli (38) skull height: the two pointers of the slide gauge are placed basally on the basis of the skull and dorsally on the highest elevation of the sagittal crest (39) skull height without the sagittal crest (40) height of the occipital triangle: from Akrokranion to Basion B. JAW MEASUREMENTS (1) total length of the lower jaw: from the Processus condyloideus to the Infradentale (7) the length of the jaw measured from the alveoli of M1 to the canine (9) the length of the jaw measured from the alveoli of M1 to the P2 (9a) the length of the jaw measured from the alveoli of M1 to the P3 (18) height measured from the basal point of the Processus angularis to the Coronion (19) the height of the lower jaw measured at the after M1 (20) the height of the lower jaw measured before the P3 C. INDIVIDUAL TEETH (cl) stands for crown length (cw) stands for crown width Super-and subscripts refer to the upper-and lower-jaws respectively. The?-symbol in graphs refers to identified teeth with lacking measurements or to identified teeth with unknown age category. 7

12 3.3.2 Postcranial measurements (GL) stands for Größe Länge : the total length of the element. (Bp) stands for Größe Breite proximal : the largest proximal width of the element. (Kd) stands for Kleinste Breite der Diaphyse : the smallest width of the diaphysis. (Bd) stands for Größe Breite distal : the largest distal width of the element Age determination The age classes of the cranial elements are based on Stiner (2004) using tooth eruption and wear: every wear stage corresponds with a code. The codes themselves are grouped in three age classes: juvenile, prime adult and old adult (fig 3.2). This ageing method was used on all separate teeth, but was also applied to the teeth of skulls and jaws. When several teeth were present, the highest code was used to assign the age-class. Sometimes identified teeth could not be assigned to an age classes, for example when the tooth-surface is broken. For this reason the total NISP in the inventory tables (section 4.1, tables 4.1 to 4.10) does sometimes not equal the sum of the NISP age class. Fig. 3.2: The wear stages and age classes, STINER (2004). d: deciduous tooth, : erupting permanent tooth, III: just erupted permanent tooth, IV-IX: erupted permanent teeth For the postcranial elements we cannot work with the table of Stiner. Therefore we classify the different elements on basis of the epiphyseal fusion of the bones and determine whether the bones belong to juveniles, sub-adults or adults (see list below). The age category for cranial remains and postcranial remains do not correspond entirely, but the postcranial ageing was mainly done to separate the juveniles; the cranial ageing is more precise. Juvenile: no fusion and small size Sub-adult: growth almost finished, but no fusion Adult: growth finished, fusion complete 8

13 3.3.4 Weathering and gnawing traces Behrensmeyer (1978) described six bone weathering stages to provide a basis for descriptive comparison. The stages are numbered from zero to five: stage 0: bones do not show signs of cracking or flaking; bones are still greasy and marrow cavities contain tissue, skin and muscle/ligament may cover part of the bone surface. stage 1: bones show cracking parallel to the fiber structure, articular surfaces may show mosaic cracking. Fat, skin or other tissues may still be present. stage 2: concentric thin layers of bone show flaking; long thin flakes are common in the initial part of this stage, followed by more extensive flaking. Remnants of ligaments, cartilage and skin may still be present. stage 3: bones have a surface characterized by patches of rough, homogeneously weathered compact bone, resulting in a fibrous texture. Tissue is rarely present. stage 4: the bone surface is coarsely fibrous and rough in texture; large and small splinters occur and may fall away from the bone when it is moved. stage 5: bones fall apart in situ, with large splinters lying around what remains of the whole, which is fragile and easily broken by moving. Original bone shape may be difficult to determine. We noted the weathering stage of each skeletal element according to these stages of Behrensmeyer. We also noted all possible traces e.g. gnawing traces and indicated on which part of the skeletal element Database Finally all data are noted in worksheets in MS Excel and lead to a database of the fossil cave hyena of Goyet, Walsin and Hastière. A separate database was made for the recent material. Based on the fossil database, inventory tables were made (section 4.1). Elements are first put in the cranial or postcranial category, then according to their age class. Then the NISP and MNIe are defined: NISP: Number of Identified Specimens; refers to the actual number of elements that were found. MNIe: Minimum Number of Individuals per skeletal Element: refers to the minimum number of individuals that can be counted from each skeletal element mainly based on counting acceptable left and right specimens. MNI: Minimum Number of Individuals: refers to the least amount of individuals in a site counting all the MNIe s. 9

14 3.3.6 Comparing fossil and recent spotted hyena The comparative study is limited to the study of the cranial material as this was the only recent material measured. We also limited ourselves to compare the adult material. We did statistical tests on the crown length and crown width of the fossil teeth to look for significant differences between the caves. For the recent material we also used a skull measurement, namely the total skull length, to see if there are significant differences present. Also the mean and standard deviation for the different teeth-types were calculated and given in mm. For these calculations we left out teeth represented only by single specimens. The number of teeth used for the calculation is noted as n. For the statistical processing (section 4.6) we use the SAS-programme, version 9_2 for windows, SAS Institute Inc., Cary, NC, USA (www. sas.com). As we have several groups to compare, we execute some ANOVA s and Tukey-Kramer s as Post hoc tests(section 4.6). 10

15 4.1 Inventory of the fossil material 4. RESULTS Tables 4.1, 4.2 and 4.3 display the inventory of the three caves (Goyet, Walsin and Hastière). Tables 4.4 until 4.10 give the inventory per cave horizon of Goyet and Hastière. NISP per skeletal element per cave The % NISP per skeletal elements of the different caves is given in fig It summarises the inventories given in tables By looking at the graphs we see that in all caves, the teeth have the highest percentage: about 54% in Goyet, 63% in Walsin and about 51% in Hastière. Because the teeth count for half (or more) of the total %NISP, it is useful to make other graphs excluding them (fig. 4.2). Now we can see that in all caves the lower jaw is the most present element with percentages of 19% (Goyet), 26% (Hastière) and 33% (Walsin). Goyet has the greatest diversity of skeletal elements (15). Besides the lower jaw, there are two more elements with a percentage above 10%: the vertebrae and the metacarpals. Then there are four elements with a percentage ranging between 5-10%: the upper jaw, the carpals/tarsals, the metatarsals and the phalanges. The other eight elements have a percentage <5%. (Fig. 4.2 A) In Walsin eight different skeletal elements were found. Two elements have, just like the lower jaw, percentages above 10%, namely the metacarpals and the phalanges. The radius and the metatarsals fall in the range of 5-10%. All the other elements present have a range <5%. (Fig. 4.2 B) Hastière is characterized by a variety of twelve different skeletal elements. Here the upper jaw and the vertebrae have also percentages of more than 10%. The ulna, metacarpals, carpals/tarsals, metatarsals and phalanges have percentages in the range of 5-10%. The other elements have a percentage < 5%. (Fig. 4.2 C) NISP per skeletal element per horizon Walsin has only one horizon, so only the horizons of Goyet and Hastière are mentioned here. Several graphs were made for each cave: with the teeth (fig. 4.3) and without the teeth (fig. 4.4 and 4.5). On figure 4.3 we can see again the dominance of the teeth over the other skeletal elements. Looking at the four horizons in Goyet (fig. 4.4), we can see that the third layer has the highest variety with a total of thirteen different skeletal elements. Horizon 4 follows with eleven different skeletal elements. In horizon 2, seven different skeletal elements were found and in horizon 1 only four. These two layers with the lowest variety have the most indeterminable elements, with percentages of 16% and 20%. In Hastière, horizon 2 has the highest diversity with a total of eleven different skeletal elements. Horizon 1 has seven and horizon 5 has four different skeletal elements. The most indeterminable elements were found in horizon 1. 11

16 GOYET all levels Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.1: Inventory of the skeletal elements found in Goyet (All horizons). Grey fields relate to elements of which the age could not be established. 12

17 Walsin Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 0 axis 0 vertebrae 0 humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus 0 calcaneum 0 carpalia/tarsalia 0 metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.2: Inventory of the skeletal elements found in Walsin. Grey fields relate to elements of which the age could not be established. 13

18 Hastière (All horizons) Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.3: Inventory of the skeletal elements found in Hastière (All horizons). Grey fields relate to elements of which the age could not be established. 14

19 GOYET horizon 1 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.4: Inventory of the skeletal elements of horizon 1, Goyet. Grey fields relate to elements of which the age could not be established. 15

20 GOYET horizon 2 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.5: Inventory of the skeletal elements found in horizon 2, Goyet. Grey fields relate to elements of which the age could not be established. 16

21 GOYET horizon 3 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.6: Inventory of the skeletal elements of horizon 3, Goyet. Grey fields relate to elements of which the age could not be established. 17

22 GOYET horizon 4 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.7: Inventory of the skeletal elements found in horizon4, Goyet. Grey fields relate to elements of which the age could not be established. 18

23 Hastière horizon 1 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP skull upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.8: Inventory of the skeletal elements found in horizon 1, Hastière. Grey fields relate to elements of which the age could not be established. 19

24 Hastière horizon 2 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP schedel upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.9: Inventory of the skeletal elements found in horizon 2, Hastière. Grey fields relate to elements of which the age could not be established. 20

25 Hastière horizon 5 Crocuta crocuta CRANIAL NISP juv. MNIe juv. NISP pr. ad. MNIe pr. ad. NISP o.ad. MNIe o.ad. Total NISP Total MNIe % NISP schedel upper jaw lower jaw dm 3 max dm 3 mand dm 4 max dm 4 mand I1 max I1 mand I2 max I2 mand I3 max I3 mand C max C mand P1 max P2 max P3 max P4 max P2 mand P3 mand P4 mand M1 mand teeth ind POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas axis vertebrae humerus ulna radius metacarpus metacarpus metacarpus metacarpus femur patella tibia fibula astragalus calcaneum carpalia/tarsalia metatarsus metatarsus metatarsus metatarsus phalanx phalanx phalanx indeterminable Total NISP Total MNI Table 4.10: Inventory of the skeletal elements found in horizon 5, Hastière. Grey fields relate to elements of which the age could not be established. 21

26 80 % NISP per skeletal element Goyet 60 % NISP A Skeletal element 80 % NISP per skeletal element Walsin 60 % NISP B Skeletal element 80 % NISP per skeletal element Hastière 60 % NISP C Skeletal element Fig. 4.1: The percentage of NISP per skeletal elements per caves. A. Goyet, B. Walsin, C. Hastière. 22

27 % NISP % NISP per skeletal element without teeth Goyet A Skeletal element % NISP % NISP per skeletal element without teeth Walsin B Skeletal element % NISP % NISP per skeletal element without teeth Hastière C Skeletal element Fig. 4.2: The percentage of NISP per skeletal elements without the teeth per caves: A. Goyet, B. Walsin, C. Hastière. 23

28 A 80 % NISP per skeletal element per horizon Goyet 60 % NISP horizon 1 horizon 2 horizon 3 horizon 4 0 Skeletal element B 80 % NISP per skeletal element per horizon Hastière 60 % NISP horizon 1 horizon 2 horizon 5 0 Skeletal element Fig. 4.3: The percentage NISP per skeletal elements per cave horizon: A. Goyet, B. Hastière 24

29 A C % NISP without teeth per skeletal element Goyet, horizon 1 % NISP without teeth per skeletal element Goyet, horizon 3 % NISP % NISP Skeletal element Skeletal element B D % NISP without teeth per skeletal element Goyet, horizon 2 % NISP without teeth per skeletal element Goyet, horizon 4 % NISP % NISP Skeletal element Skeletal element Fig. 4.4: The percentage NISP without teeth - Goyet per horizon: A. Horizon 1, B. Horizon 2, C. Horizon 3, D. Horizon 4 23

30 15 % NISP per skeletal element without teeth Hastière, horizon 1 % NISP A Skeletal element 15 % NISP per skeletal element without teeth Hastière, horizon 2 % NISP B Skeletal element 15 % NISP per skeletal element without teeth Hastière, horizon 5 % NISP C Skeletal element Fig. 4.5: The percentage NISP of Hastière per horizon: A. Horizon 1, B. Horizon 2, C. Horizon 5 26

31 4.2 Description of the fossil material Cranial A. CRANIUM Three skulls were measured: one from Goyet, one from Hastière and one skull that was found in the North Sea (table 4.11). The skull from Goyet was not complete and several measurements are lacking. Site Goyet Hastière North sea Horizon?? - Number DP 4017 (veri 00-40) 2197 (veri 00-44) - (1) (2) (8) (12) (15) (25) (27) (28) (29) (34) (35) (36) (38) (39) (40) Table 4.11: The measurements of the fossil skulls. Photo 4.1: Skull with teeth (P2, P3 and P4) from Hastière specimen 2197 (veri 00-44). 27

32 B. UPPER JAW In Goyet a total of fourteen upper jaws were found: two in horizon 2, four in horizon 3 and eight in horizon 4 (tables ). In Walsin, only one upper jaw was found and it belonged to a prime adult (table 4.2). Hastière contained a total of ten upper jaws: five in horizon one, four in horizon 2 and one in horizon 5 (tables ). Except for the dentition, no measurements could be made as all upper jaws were broken. Photo 4.2: Upper jaw with P3 and P4 from Goyet (horizon 4) specimen C. LOWER JAW Goyet contained a total of thirty-six lower jaws. Most of the jaws were found in horizon 4, namely 22 specimens. Horizon 3 had eight, horizon 2 had four and horizon 1 had two lower jaws (tables ). Walsin harboured nine mandibles. In Hastière 25 lower jaws were found: seven in horizon 1, 17 in horizon 2 and one in horizon 5. Table 4.12 gives the available measurements of the adult jaws. Site Horizon Plateau Number ID DP (1) (7) (9) (9a) (18) (19) (20) Goyet Goyet Goyet Goyet Goyet Goyet Goyet Goyet Goyet Walsin Walsin Walsin

33 Hastière Hastière Hastière Hastière Hastière Table 4.12: The measurements of the fossil lower jaws. Photo 4.3: Lower jaw with C, P3, P4 and M1 of a prime adult from Goyet (horizon 4) specimen Photo 4.4: Lower jaw with di2, di3, dc, dm2, dm3 and dm4 of a juvenile from Goyet (horizon 4) specimen a 29

34 D. TEETH Deciduous teeth Not many isolated deciduous teeth were found, the total number in all caves amounts to five. Three were found in Goyet: one dm3 and two dm4. From the latter, one tooth could not be identified as belonging to the upper or lower jaw as it was broken; it was counted in table 4.1 as an indeterminable tooth. In Hastière one dm3 and one dm4 was found, whilst in Walsin no isolated deciduous teeth were found. Table 4.13 gives all the details about these five teeth. Cave Horizon Plateau Number Position Crown Crown ID Plateau Tooth DP in jaw length width Goyet dm 3 upper left Goyet dm Goyet dm 4 lower left Hastière b 99 dm 3 upper left Hastière dm 4 lower left Table 4.13: The isolated deciduous teeth Table 4.14 gives the mean and standard deviation of the different deciduous teeth. For these calculations not only the isolated deciduous teeth were used but also the teeth from the upper and lower jaws. Cave Goyet Dental type n Mean Std dev dm 2 cl dm 2 cw dm 3 cl dm 3 cw dm 3 cl dm 3 cw dm 4 cl dm 4 cw Hastière dm 4 cl dm 4 cw Table 4.14: All deciduous teeth from Goyet and Hastière. 30

35 Incisors In total 49 isolated incisors were found and Goyet has the highest number of incisors with a total number of 22. Of this amount there were three I1 s, one I2 and 18 I3 s. Walsin had a total of eight incisors with one I1, two I2 s and five I3 s. In Hastière 19 incisors were found: one I1, three I2 s and six I3 s (tables ). Table 4.15 gives the measurements of all incisors per cave. Cave Dental type n Mean Std dev I 1 cl Goyet Walsin Hastière I 1 cw I 3 cl I 3 cw I 1 cl I 1 cw I 3 cl I 3 cw I 2 cl I 2 cw I 3 cl I 3 cw I 3 cl I 3 cw Table 4.15: The incisors of the three caves. Canines In Goyet 54 upper and 23 lower canines were found, Walsin had only five upper and two lower canines and Hastière had 12 upper canines and 14 lower canines (tables ). Figure 4.6 gives the percentages of the upper and lower canines found in Goyet per crown length classes. Both isolated teeth and teeth still in the jaw were used here. We can see that most canines, both upper and lower, fall in a range of 15 and 18 millimetres suggesting an unimodal distribution. We have not made such a figure for the canines found in Walsin because of the few finds. Instead we give a table with the measurement of the canines of Walsin (table 4.16). Figure 4.7 gives the percentages per crown length classes of the canines found in Hastière. Again both isolated teeth and teeth still in the jaws were used for the construction of this graph. Here most upper canines fall in the range of 16 to 19 mm and represent an unimodal distribution. The lower canines have a somewhat lower range between 15 and 17 mm and present also an unimodal distribution. Table 4.17 gives the mean and standard deviation of the crown lengths and widths of all measurable canines found in the three caves, using both isolated canines and these of the jaws for the calculation. 31

36 Cave Number DP Number ID plateau position Crown length Crown width Walsin upper left Walsin upper left Walsin upper right Walsin upper right Walsin upper right Walsin lower left Walsin lower right Walsin lower left mand Walsin lower right mand Table 4.16: The canines of Walsin and their measurements. Cave Dental type n Mean Std dev C max cl Goyet Walsin Hastière C max cw C mand cl C mand cw C max cl C max cw C mand cl C mand cw C max cl C max cw C mand cl C mand cw Table 4.17: The canines of the three caves. 32

37 Percentage of C per crown length class Goyet Percentage (in%) ? C max C mand Crown length in classes (in mm) Figure 4.6: The frequency distributions of the crown length of the upper and lower canines found in Goyet. Percentage of C per crown length class Hastière Percentage (in %) ? C max C mand Crown length in classes (in mm) Figure 4.7: The frequency distributions of the crown length of the upper and lower canines found in Hastière. Premolars In Goyet, not less than 75 isolated premolars were identified. Walsin had only 14 premolars, Hastière did somewhat better with 34 isolated premolars (tables ). Table 4.18 gives the dental type, the mean and standard deviation of the crown lengths and crown widths of the four different premolars. Like in the previous tables, isolated and teeth still in the jaw were used. For the upper and lower P3 s and the lower P4 s graphs (fig ) give again the frequency distribution of the crown length. We used the upper and lower P3 and lower P4 because of these at least 30 teeth were present; this was not the case for the upper and lower P2 s and the upper P4 s. In fig. 4.9 there is a tendency for a unimodal curve in the upper P3 crown length, for the lower P3-curve the trend is less clear. 33

38 Dental type n Mean Std dev P 2 cl P 2 cw P 2 cl P 2 cw P 3 cl P 3 cw P 3 cl P 3 cw P 4 cl P 4 cw P 4 cl P 4 cw Table 4.18: The premolars of Goyet. 50 Percentage of P3 per crown length class Goyet Percentage (in %) P3 max P3 mand ? Crown length in classes (in mm) Fig. 4.8: Frequency distribution of the crown length of the upper and lower P3 s. Percentage (in %) Percentage of lower P4 per crown length class Goyet Crown length in classes (in mm) Fig. 4.9: Frequency distribution of the crown length of the lower P4 s. 34

39 For Walsin, table 4.19 gives the mean and standard deviation; as the material is not very numerous, no figures like 4.8 or 4.9 were made. Table 4.20 displays the measurements of Hastière; again no graphs were made. Dental type n Mean Std dev Dental type n Mean Std dev P 2 cl P 2 cw P 3 cl P 3 cw P 3 cl P 3 cw P 4 cl P 4 cw P 4 cl P 4 cw Table 4.19: The premolars of Walsin. P 2 cl P 2 cw P 2 cl P 2 cw P 3 cl P 3 cw P 3 cl P 3 cw P 4 cl P 4 cw P 4 cl P 4 cw Table 4.20: The premolars of Hastière. Molars Goyet has the most isolated molars, namely 25. Walsin has only five isolated molars and Hastière has only eight isolated molars. For all the molars present, table 4.21 gives the calculated means and standard deviations. Cave Horizon Dental type n Mean Std dev horizon 1 M1 cl M1 cw Goyet horizon 2 M1 cl M1 cw horizon 3 M1 cl M1 cw horizon 4 M1 cl M1 cw Walsin M1 cl M1 cw Hastière horizon 1 M1 cl M1 cw horizon 2 M1 cl M1 cw Table 4.21: The molars of all the caves. 35

40 For the molars, in contrast to the premolars, we did make graphs with the amount of teeth per crown length classes, even if Walsin and Hastière do not have each a minimum of 30 molars (fig. 4.10). We did this because we wanted to compare at least some type of the teeth over the three caves. The molars in Goyet have the highest amount in the class, the class has the second most. In this graph we can see a sort of bimodal pattern (fig. 4.10B). Walsin shows no such pattern; it has three classes with the same amount of teeth: 30-31, and We see a unimodal pattern here but we do want to remind that only 8 molars were found in the cave (fig. 4.10C). Hastière has also a unimodal pattern and here the highest number is found in the class (fig. 4.10D). If we look at all the caves together, the highest number of teeth fall between the range of mm. 36

41 A C Percentage of M1 per cl class all caves Percentage of M1 per cl class Walsin Percentage (in %) Percentage (in %) Crown length in classes (in mm) Crown length in classes (in mm) B D Percentage of M1 per cl class Goyet Percentage of M1 per cl class Hastière Percentage (in %) Percentage (in %) Crown length in classes (in mm) Crown length in classes (in mm) Fig. 4.10: The amount of M1 s per crown length classes. A. All caves, B. Goyet, C. Walsin and D. Hastière. 37

42 4.2.2 Postcranial A. VERTEBRAE Here we only distinguished between the atlas, axis and the other vertebrae. For the calculation of the %NISP per skeletal element we counted all vertebrae together. Only the Goyet cave and the Hastière cave harboured vertebrae: 20 specimens in Goyet and 15 in Hastière, good for 5.0% and 7.7% of the total NISP (tables 4.1 and 4.3, fig.4.1). For the frequency of vertebrae per horizon, we refer to figure 4.3. B. HUMERUS Eight fragments of humeri were found, six pieces in Goyet (1.5% of total NISP) and two in Hastière (1.0% of total NISP) (tables 4.1 and 4.3, fig. 4.1). As there were only fragments of the humeri, only two measurements could be made, we refer to them in table Site Horizon Number DP Plateau ID (Bd) (Kd) Goyet Goyet Table 4.22: Measurements of the humeri. Photo 4.5: Humerus: left distal from Goyet (horizon 4) specimen C. ULNA In the three caves, a total of 18 ulnae were found, with nine in Goyet (2.3% of total NISP), one in Walsin (1.4% of total NISP)and eight in Hastière (4.1% of total NISP) (tables , fig. 4.1). Again, like the humeri, most elements were too much broken and/or incomplete to get good measurements. Measurements from an ulna of Goyet are listed below in table 4.23: Site Horizon Number DP Plateau ID (GL) (Kd) (Bd) Goyet Table 4.23: Measurements of the ulna. 38

43 D. RADIUS In total eight radii were found (tables , fig. 4.1). Table 4.24 gives the available measurements of these radii. Site Horizon Number DP Plateau ID (GL) (Bp) (Kd) (Bd) Goyet Goyet Walsin Walsin Hastière A Table 4.24: Measurements of the radii. Photo 4.6: Radius (left) with gnawing traces from Goyet (horizon 2) specimen E. CARPALIA AND TARSALIA For these skeletal elements we followed the identification as noted by Dupont (unpublished notes). We grouped the different carpals and tarsals together. This gives us a total of 18 elements found in Goyet and Hastière, as none were found in Walsin (tables ). F. METACARPALS The counts per cave and per horizon of the different metacarpals are shown in tables 4.1 to 4.10 (fig 4.1). See table 4.25 for the measurements. G. FEMUR One femur was found, located in the third horizon of Goyet. It is a right diaphysis of an adult. It shows a weathering stage 1 and it does not bear any gnawing traces. The only available measurement is the smallest width of the diaphysis (Kd): Kd = mm. 39

44 Site Horizon Number DP Plateau ID Element (GL) (Bp) (Kd) (Bd) Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Hastière i MC Hastière c MC Hastière f MC Walsin MC Walsin MC Walsin MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Goyet MC Hastière c MC Hastière b MC Walsin MC Goyet MC Goyet MC Goyet MC Goyet MC Hastière e MC Hastière b MC Hastière MC Hastière b MC Walsin MC Table 4.25: Measurements of the metacarpals. 40

45 H. PATELLA The only patella found was located in the third horizon of Goyet. It is a right one and its edges are at one side worn off. It probably belonged to an adult individual but no measurements were made. I. TIBIA A total of nine tibias were found, table 4.26 gives an overview of the available measurements. Site Niveau Number DP plateau ID (GL) (Bp) (Kd) (Bd) Goyet Goyet Goyet Walsin Hastière Hastière Table 4.26: Measurements of the tibiae. J. FIBULA One fibula was found in the third horizon of Goyet. It is a right diaphysis of a juvenile. It shows weathering stage 1 and no gnawing traces. The only measurement possible is the smallest width of the diaphysis (Kd): Kd= K. METATARSALS The different metatarsals are shown in tables 4.1 to 4.10 for the counts per cave and per horizon. Table 4.27 gives the available measurements. L. PHALANGES As for the metacarpals and metatarsals we first differentiated between the different phalanges; their counts can be seen in tables 4.1 to 4.3 for each cave and in tables 4.4 to 4.10 for each horizon. For the three caves, a total of 21 phalanges were found: 12 of them were situated in Goyet (3.0% of the total NISP), three in Walsin (4.1%) and six in Hastière (3.1%) (fig. 4.1). The phalanges were not measured. 41

46 Site Horizon Number DP Plateau ID Element (GL) (Bp) (Kd) (Bd) Goyet MT Goyet MT Hastière d MT Goyet MT Goyet MT Goyet MT Goyet bis 29 MT Goyet MT Walsin MT Goyet MT Goyet MT Goyet MT Hastière e MT Goyet MT Goyet MT Goyet MT Goyet MT Hastière d MT Hastière d MT Walsin MT Table 4.27: Measurements of the metatarsals. 42

47 4.3 Age determination We made some frequency distributions on the wear-stages (STINER, 2004) of the fossil M1 s per cave (fig ). Goyet and Hastière have three age categories: juveniles ( germ to III), prime adults (IV to VII) and old adults (VIII and IX). In Walsin we find prime adults and old adults. We also made frequency distributions of the age classes (according to Stiner) of the fossil upper and lower jaws (fig. 4.12). Also here, Goyet and Hastière have juveniles, prime adults and old adults; whilst Walsin has only prime and old adults. For the age of the postcranial elements we refer to the inventory tables (section 4.1) and to the tables made in section Percentage (in %) Percentages of M1 per wear stages (Stiner) Goyet germ I II III IV V VI VII VIII IX? A B C Percentage (in %) Percentage (in %) Wear stages (according to stiner) Percentages of M1 per wear stages (Stiner) Walsin germ I II III IV V VI VII VIII IX? Wear stages (according to stiner) Percentages of M1 per wear stages (Stiner) Hastière germ I II III IV V VI VII VIII IX? Wear stages (according to Stiner) Fig. 4.11: The frequency distributions of the teeth per wear stages (Stiner). A. Goyet, B. Walsin, C. Hastière 43

48 Percentage of upper jaws per age class Percentage (in %) Juvenile Prime adult Old adult? Goyet Walsin Hastière A Age classes (according to stiner) Percentage of lower jaws per age class 100 Percentage (in %) juvenile prime adult old adult? Goyet Walsin Hastière B Age classes (according to Stiner) Fig. 4.12: The percentages of the jaws per age classes per caves. A. Upper jaws, B. Lower jaws For the recent material we made a frequency distribution of the wear stages of M1 per sex (fig. 4.13). There are juvenile males but no juvenile females. The specimens with unknown gender all belong to the prime adult age class or to the old adult age class. Percentage (in %) Percentages of M1 per wear stages (Stiner) Recent material germ I II III IV V VI VII VIII IX? Wear stages (according to stiner) Fig. 4.13: Frequency distribution of the wear stages of M1 of the recent material. 44

49 4.4 Trace fossils Coprolites In table 4.28 we give an overview of the coprolites found per cave and per horizon. Cave n coprolites Goyet - all horizons 4 Goyet - horizon 1 - Goyet - horizon 2 - Goyet - horizon 3 - Goyet - horizon 4 4 Walsin - Hastière - all horizons 31 Hastière - horizon 1 5 Hastière - horizon 2 25 Hastière - horizon 5 1 Table 4.28: Coprolites Other traces The different skeletal elements of the fossil material were checked for traces. More specific we looked if there were gnawing marks and/or cut-traces, marks of ochre or if there were any other interesting traces. We only found gnawing marks and table 4.29 gives all the detailed information about these skeletal elements. The frequency distribution of the skeletal elements with gnawing marks based on the NISP per horizon is shown in fig. 4.14A. We see that the second horizon of Hastière has the highest percentage of gnawed elements. Most elements that were gnawed belonged to juveniles and sub-adults (fig. 4.14B). 45

50 Site Horizon Id Plateau Element Position Age Broken Weathering Gnawing traces Goyet bone fragment - - x 1 proximal and distal Goyet MT3 left juvenile x 1 proximal Goyet MT4 left juvenile x 1 distal Goyet radius left proximal and distal Goyet MC2 right adult - 1 proximal and distal Goyet MC2 left sub-adult - 1 proximal and distal Goyet MC5 right sub-adult x 1 proximal Goyet humerus right adult x 1 distal Goyet humerus right adult x 1 distal Goyet ulna left juvenile x 1 proximal Goyet ulna left adult x 1 proximal Goyet MC2 right juvenile - 1 proximal and distal Goyet MC5 right sub-adult - 1 proximal en diaphyse Goyet MC5 right sub-adult - 1 proximal and distal Goyet radius right proximal and distal Walsin MC5 left juvenile - 1 distal Walsin radius right adult - 1 diaphyse Hastière ulna proximal right juvenile x 1 distal Hastière 2 2h 99 MT2 left juvenile - 1 diaphyse Hastière 2 5c 99 MC4 distal right juvenile x 1 proximal en diaphyse Hastière 2 4b 99 MT4 proximal right sub-adult - 1 distal Hastière 2 1e 99 MT4 left sub-adult - 1 distal Hastière 2 7b 99 MC5 left sub-adult - 1 proximal and distal Hastière 2 4d 99 MT5 proximal right juvenile - 1 distal Hastière 2 2c 99 humerus proximal - sub-adult x 1 whole piece Hastière 2 1a 99 radius proximal right - x 1 proximal and distal Hastière 2 1d 99 tibia distal right juvenile x 1 distal Hastière 2 1c 99 ulna proximal right sub-adult x 1 distal Table 4.29: The skeletal elements with gnawing traces. 46

51 Percentage (in %) Percentage skeletal elements with gnaw traces per cave and per horizon Percentage (in %) Skeletal elements with gnawing traces: percentages per age class Juvenile sub adult adult? Age classes Fig. 4.14: A. Percentage skeletal elements with gnaw-traces, B. Skeletal elements with gnawing traces: percentages per age class. 4.5 Weathering All skeletal elements had weathering stage 1, except for one tibia that was found in the fourth horizon of Goyet. This tibia has a weathering stage 2; photograph 4.7 shows this tibia in more detail. Photo 4.8: Tibia (left) from Goyet (horizon 4) specimen