Fossil Cave Hyena of Goyet, Walsin and Hastière (Belgium): Osteometry and Taphonomy

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FACULTY OF SCIENCES Palaeontology Department Biology Department

Academic year 2009-2010

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)

Table of contents

1. INTRODUCTION 1

1.1. General introduction 1 1.2. The spotted hyena - Crocuta crocuta 1 1.2.1. Recent spotted hyena 1 1.2.2. Fossil spotted hyena or cave hyena 2

2. AIMS 2

3. MATERIAL AND METHODS 3

3.1 Description of the sites 3 3.2 Description and organisation of the material 6 3.3 Methods and measurements 6 3.3.1 Cranial measurements 6 3.3.2 Postcranial measurements 8 3.3.3 Age determination 8 3.3.4 Weathering and gnawing traces 9 3.3.5 Database 9 3.3.6 Comparing fossil and recent spotted hyena 10

4. RESULTS 11

4.1. Inventory of the fossil material 11 4.2. Description of the fossil material 27 4.2.1. Cranial 27 4.2.2. Postcranial 38 4.3. Age determination 43 4.4. Trace fossils 45 4.4.1. Coprolites 45 4.4.2. Other traces 45 4.5. Weathering 47 4.6. Inventory of the recent material 48 4.7. Description of the recent material 50 4.7.1. Cranial 50 4.8. The fossil and recent spotted hyena 61

5. DISCUSSION 66

5.1. Taphonomy 66 5.2. Age determination – Use as communal dens? 68 5.3. Gnawing traces and cannibalism 70 5.4. Fossil and recent spotted hyena 71

6. CONCLUSIONS 72

7. DUTCH SUMMARY 73

8. THANKS 75

9. REFERENCES 76

ANNEXES 79

Alphabetical List of abbreviations 79

Statistical tests CD-Rom

Primary data CD-Rom

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 1872. 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

1.2.1 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.

1.2.2 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.

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 1869. 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 (110-120 m) (GERMONPRÉ, 1996, 2001, 2004).

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 12 620 +/- 90

horse 12 770 +/- 90

27 230 +/- 260 cave hyena 35 000 +/- 400

cave bear 38 770+1180-1030

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).

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 1905-1906, 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.

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 (1902-1912), 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.

3.3.1 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

(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.

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.

3.3.3 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

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.

3.3.5 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.

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).

4. RESULTS

4.1 Inventory of the fossil material

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. 4.1. It summarises the inventories given in tables 4.1-4.3. 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.

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 0 0 1 1 0 0 1 1 0.25 upper jaw 2 1 9 5 2 1 14 7 3.48 lower jaw 6 6 19 10 3 3 36 19 8.96 dm 3 max 1 1 0 0 0 0 1 1 0.25 dm 3 mand000 000000.00 dm 4 max 0 0 0 0 0 0 0 0 0.00 dm 4 mand110 000110.25 I1 max 0 0 2 2 0 0 3 2 0.75 I1 mand000 000000.00 I2 max 1 1 0 0 0 0 1 1 0.25 I2 mand000 000000.00 I3 max 1 1 8 5 5 4 16 10 3.98 I3 mand001 100210.50 C max 54 28 13.43 C mand 23 17 5.72 P1 max 0 0 0 0 0 0 0 0 0.00 P2 max 0 0 0 0 0 0 0 0 0.00 P3 max 5 4 8 6 2 2 16 12 3.98 P4 max 0 0 4 3 2 2 7 5 1.74 P2 mand 4 3 6 4 0 0 10 7 2.49 P3 mand 3 2 17 8 0 0 22 10 5.47 P4 mand 3 3 14 8 1 1 20 12 4.98 M1 mand 3 2 15 11 6 5 25 18 6.22 teeth ind. 13 3.23 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 20.50 axis 10.25 vertebrae 17 4.23 humerus 0 0 1 1 5 4 6 5 1.49 ulna423 222962.24 radius11 21320.75 metacarpus 2211 143751.74 metacarpus 3000 032320.75 metacarpus 4001 152631.49 metacarpus 5002 244661.49 femur 0 0 0 0 1 1 1 1 0.25 patella 10.25 tibia 0 0 0 0 4 2 4 2 1.00 fibula 1 1 0 0 0 0 1 1 0.25 astragalus 20.50 calcaneum 30.75 carpalia/tarsalia 81.99 metatarsus 2001 111220.50 metatarsus 3113 221641.49 metatarsus 4221 111441.00 metatarsus 5331 100441.00 phalanx 10011961072.49 phalanx 2000011110.25 phalanx 3000011110.25 indeterminable 27 6.72

Total NISP 44 119 66 400 Total MNI 6 11 6 34

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.

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 0 0 1 1 0 0 1 1 1.37 lower jaw 0 0 7 4 2 1 9 5 12.33 dm 3 max 0 0 0 0 0 0 0 0 0.00 dm 3 mand000 000000.00 dm 4 max 0 0 0 0 0 0 0 0 0.00 dm 4 mand000 000000.00 I1 max 0 0 1 1 0 0 1 1 1.37 I1 mand000 000000.00 I2 max 1 1 0 0 0 0 1 1 1.37 I2 mand000 011111.37 I3 max 1 1 4 2 0 0 5 3 6.85 I3 mand000 000000.00 C max 5 3 6.85 C mand 2 1 2.74 P1 max 0 0 0 0 0 0 0 0 0.00 P2 max 0 0 0 0 0 0 0 0 0.00 P3 max 3 3 1 1 0 0 4 4 5.48 P4 max 0 0 2 1 0 0 2 1 2.74 P2 mand000 000000.00 P3 mand003 200324.11 P4 mand212 211546.85 M1 mand002 232546.85 teeth ind. 12 16.44 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 0 0 0 0 0 0 0 0 0.00 ulna 0 0 0 0 1 1 1 1 1.37 radius 0 0 1 1 1 1 2 2 2.74 metacarpus 2111 111334.11 metacarpus 3000 000000.00 metacarpus 4110 000111.37 metacarpus 5110 000111.37 femur 0 0 0 0 0 0 0 0 0.00 patella000 000000.00 tibia 0 0 1 1 0 0 1 1 1.37 fibula 0 0 0 0 0 0 0 0 0.00 astragalus 0 calcaneum 0 carpalia/tarsalia 0 metatarsus 2000 000000.00 metatarsus 3110 000111.37 metatarsus 4000 000000.00 metatarsus 5110 000111.37 phalanx 1000033334.11 phalanx 2000000000.00 phalanx 3000000000.00 indeterminable 34.11

Total NISP 12 26 13 73 Total MNI 3 4 3 8

Table 4.2: Inventory of the skeletal elements found in Walsin. Grey fields relate to elements of which the age could not be established.

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 00 1 10 01 10.51 upper jaw 2 1 6 5 2 2 10 8 5.13 lower jaw 4 3 18 10 3 2 25 15 12.82 dm 3 max11 0 00 01 10.51 dm 3 mand000 000000.00 dm 4 max00 0 00 00 00.00 dm 4 mand110 000110.51 I1 max00 1 10 01 10.51 I1 mand000 000000.00 I2 max00 0 00 00 00.00 I2 mand112 200331.54 I3 max11 5 42 19 74.62 I3 mand002 222653.08 C max 12 8 6.15 C mand 14 9 7.18 P1 max00 0 00 00 00.00 P2 max11 3 20 04 32.05 P3 max00 7 40 07 43.59 P4 max11 6 40 08 54.10 P2 mand320 000321.54 P3 mand222 111653.08 P4 mand114 300542.56 M1 mand324 311864.10 teeth ind. 11 5.64 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 21.03 axis 00.00 vertebrae 13 6.67 humerus00 1 11 12 21.03 ulna423 311864.10 radius22 1 10 03 31.54 metacarpus 2 21 1 10 03 21.54 metacarpus 3 00 0 00 00 00.00 metacarpus 4 11 1 10 02 21.03 metacarpus 5 11 3 20 04 32.05 femur00 0 00 00 0.00 patella000 000000.00 tibia 32 1 10 04 32.05 fibula00 0 00 00 00.00 astragalus 10.51 calcaneum 00.00 carpalia/tarsalia 42.05 metatarsus 2 11 1 10 02 21.03 metatarsus 3 00 0 00 00 00.00 metatarsus 4 00 1 10 01 10.51 metatarsus 5 11 1 10 02 21.03 phalanx 1 00 0 05 45 42.56 phalanx 2 00 0 01 11 10.51 phalanx 3 00 0 00 00 00.00 indeterminable 31.54

Total NISP 36 75 19 195 Total MNI 3 10 4 15

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.

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 0 0 0 0 0 0 0 0 0.00 lower jaw 1 1 1 1 0 0 2 2 4.65 dm 3 max 0 0 0 0 0 0 0 0 0.00 dm 3 mand000 000000.00 dm 4 max 0 0 0 0 0 0 0 0 0.00 dm 4 mand000 000000.00 I1 max 0 0 2 2 0 0 3 2 6.98 I1 mand000 000000.00 I2 max 1 1 0 0 0 0 1 1 2.33 I2 mand000 000000.00 I3 max 0 0 0 0 0 0 0 0 0.00 I3 mand000 000000.00 C max 409.30 C mand 306.98 P1 max 0 0 0 0 0 0 0 0 0.00 P2 max 0 0 0 0 0 0 0 0 0.00 P3 max 2 1 2 1 1 1 5 3 11.63 P4 max 0 0 0 0 0 0 0 0 0.00 P2 mand000 000000.00 P3 mand003 300439.30 P4 mand002 111326.98 M1 mand113 3115511.63 teeth ind. 102.33 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 00.00 axis 00.00 vertebrae 00.00 humerus 0 0 0 0 0 0 0 0 0.00 ulna000 000000.00 radius000 000000.00 metacarpus 2110 000112.33 metacarpus 3000 011112.33 metacarpus 4000 000000.00 metacarpus 5000 000000.00 femur 0 0 0 0 0 0 0 0 0.00 patella 00.00 tibia 0 0 0 0 0 0 0 0 0.00 fibula 0 0 0 0 0 0 0 0 0.00 astragalus 00.00 calcaneum 12.33 carpalia/tarsalia 00.00 metatarsus 2000 000000.00 metatarsus 3000 000000.00 metatarsus 4000 000000.00 metatarsus 5000 000000.00 phalanx 1000022224.65 phalanx 2000000000.00 phalanx 3000000000.00 indeterminable 716.28

Total NISP 6 13 6 43 Total MNI 1 3 2 5

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.

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 jaw11 1 10 02 22.86 lower jaw00 2 10 04 25.71 dm 3 max00 0 00 00 00.00 dm 3 mand 0 0 0 0 0 0 0 0 0.00 dm 4 max11 0 00 01 11.43 dm 4 mand 0 0 0 0 0 0 0 0 0.00 I1 max00 0 00 00 00.00 I1 mand 0 0 0 0 0 0 0 0 0.00 I2 max00 0 00 00 00.00 I2 mand 0 0 0 0 0 0 0 0 0.00 I3 max00 1 12 23 34.29 I3 mand 0 0 0 0 0 0 1 1 1.43 C max 10 8 14.29 C mand 334.29 P1 max00 0 00 00 00.00 P2 max00 0 00 00 00.00 P3 max00 1 11 13 34.29 P4 max00 1 10 01 11.43 P2 mand 0 0 1 1 0 0 1 1 1.43 P3 mand 1 1 3 2 0 0 4 3 5.71 P4 mand 0 0 2 1 0 0 2 1 2.86 M1 mand 0 0 4 2 3 3 8 6 11.43 teeth ind. 11.43 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 00.00 axis 00.00 vertebrae 22.86 humerus00 0 00 00 00.00 ulna 0 0 0 0 0 0 0 0 0.00 radius 1 1 0 0 1 1 2 2 2.86 metacarpus 2 00 0 00 00 00.00 metacarpus 3 00 0 00 00 00.00 metacarpus 4 00 0 00 00 00.00 metacarpus 5 00 0 00 00 00.00 femur00 0 00 00 00.00 patella 00.00 tibia 00 0 00 00 00.00 fibula 0 0 0 0 0 0 0 0 0.00 astragalus 11.43 calcaneum 11.43 carpalia/tarsalia 00.00 metatarsus 2 00 0 00 00 00.00 metatarsus 3 11 0 00 01 11.43 metatarsus 4 11 0 00 01 11.43 metatarsus 5 00 0 00 00 00.00 phalanx 1 00 0 04 44 45.71 phalanx 2 00 0 00 00 00.00 phalanx 3 00 0 01 11 11.43 indeterminable 14 20.00

Total NISP 6 16 12 71 Total MNI 1 2 4 9

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.

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 jaw11 2 21 14 42.74 lower jaw11 6 41 18 65.48 dm 3 max11 0 00 01 10.68 dm 3 mand 0 0 0 0 0 0 0 0 0.00 dm 4 max00 0 00 00 00.00 dm 4 mand 1 1 0 0 0 0 1 1 0.68 I1 max00 0 00 00 00.00 I1 mand 0 0 0 0 0 0 0 0 0.00 I2 max00 0 00 00 00.00 I2 mand 0 0 0 0 0 0 0 0 0.00 I3 max00 4 32 27 54.79 I3 mand 0 0 1 1 0 0 1 1 0.68 C max 22 11 15.07 C mand 11 8 7.53 P1 max00 0 00 00 00.00 P2 max00 0 00 00 00.00 P3 max11 1 10 02 21.37 P4 max00 1 11 12 21.37 P2 mand 4 3 4 3 0 0 8 6 5.48 P3 mand 1 1 6 5 0 0 8 6 5.48 P4 mand 2 2 8 4 0 0 12 6 8.22 M1 mand 0 0 2 1 0 0 2 1 1.37 teeth ind. 11 7.53 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 10.68 axis 00.00 vertebrae 42.74 humerus00 1 14 25 33.42 ulna 1 1 2 1 1 1 4 3 2.74 radius 000.00 metacarpus 2 00 1 12 13 22.05 metacarpus 3 00 0 01 11 10.68 metacarpus 4 00 1 12 13 22.05 metacarpus 5 00 1 11 12 21.37 femur00 0 01 11 10.68 patella 10.68 tibia 00 0 01 11 10.68 fibula 1 1 0 0 0 0 1 1 0.68 astragalus 10.68 calcaneum 00.00 carpalia/tarsalia 53.42 metatarsus 2 00 1 10 01 10.68 metatarsus 3 00 3 21 14 32.74 metatarsus 4 11 0 00 01 10.68 metatarsus 5 32 0 00 03 22.05 phalanx 1 00 1 12 23 32.05 phalanx 2 00 0 00 00 00.00 phalanx 3 00 0 00 00 00.00 indeterminable 53.42

Total NISP 18 46 21 150 Total MNI 3 5 2 14

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.

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 0 0 6 3 1 1 8 4 5.80 lower jaw 10 5 10 6 2 2 22 13 15.94 dm 3 max 0 0 0 0 0 0 0 0 0.00 dm 3 mand000 000000.00 dm 4 max 0 0 0 0 0 0 0 0 0.00 dm 4 mand000 000000.00 I1 max 0 0 0 0 0 0 0 0 0.00 I1 mand000 000000.00 I2 max 0 0 0 0 0 0 0 0 0.00 I2 mand000 000000.00 I3 max 1 1 3 3 1 1 6 5 4.35 I3 mand000 000000.00 C max 18 12 13.04 C mand 654.35 P1 max 0 0 0 0 0 0 0 0 0.00 P2 max 0 0 0 0 0 0 0 0 0.00 P3 max 2 2 4 3 0 0 6 5 4.35 P4 max 0 0 3 2 1 1 4 3 2.90 P2 mand001 100110.72 P3 mand115 300644.35 P4 mand112 200332.17 M1 mand 2 1 6 5 2 2 10 8 7.25 teeth ind. 10.72 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 10.72 axis 10.72 vertebrae 11 7.97 humerus 0 0 0 0 1 1 1 1 0.72 ulna311 111533.62 radius000 011110.72 metacarpus 2110 022332.17 metacarpus 3000 011110.72 metacarpus 4000 032322.17 metacarpus 5001 133442.90 femur 0 0 0 0 0 0 0 0 0.00 patella 00.00 tibia 0 0 0 0 3 2 3 2 2.17 fibula 0 0 0 0 0 0 0 0 0.00 astragalus 00.00 calcaneum 10.72 carpalia/tarsalia 10.72 metatarsus 2000 000000.00 metatarsus 3000 011110.72 metatarsus 4110 011221.45 metatarsus 5110 000110.72 phalanx 1000011110.72 phalanx 2000011110.72 phalanx 3000000000.00 indeterminable 21.45

Total NISP 23 42 26 135 Total MNI 5 6 3 13

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.

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 jaw11 4 40 05 59.62 lower jaw00 6 31 17 413.46 dm 3 max00 0 00 00 00.00 dm 3 mand000 000000.00 dm 4 max00 0 00 00 00.00 dm 4 mand000 000000.00 I1 max00 0 00 00 00.00 I1 mand000 000000.00 I2 max00 0 00 00 00.00 I2 mand000 000000.00 I3 max00 1 11 13 25.77 I3 mand001 111223.85 C max 4 3 7.69 C mand 5 4 9.62 P1 max00 0 00 00 00.00 P2 max00 0 00 00 00.00 P3 max00 4 20 04 27.69 P4 max00 1 10 01 11.92 P2 mand 0 0 0.00 P3 mand000 011111.92 P4 mand111 100223.85 M1 mand001 100111.92 teeth ind. 35.77 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 00.00 axis 00.00 vertebrae 47.69 humerus00 0 00 00 00.00 ulna212 211549.62 radius00 0 00 00 00.00 metacarpus 2 00 0 00 00 00.00 metacarpus 3 00 0 00 00 00.00 metacarpus 4 00 0 00 00 00.00 metacarpus 5 00 0 00 00 00.00 femur 00.00 patella000 000000.00 tibia 11 0 00 01 11.92 fibula 00.00 astragalus 00.00 calcaneum 00.00 carpalia/tarsalia 00.00 metatarsus 2 00 0 00 00 00.00 metatarsus 3 00 0 00 00 00.00 metatarsus 4 00 0 00 00 00.00 metatarsus 5 00 0 00 00 00.00 phalanx 1 00 0 01 11 11.92 phalanx 2 00 0 00 00 00.00 phalanx 3 00 0 00 00 00.00 indeterminable 35.77

Total NISP 5 21 6 52 Total MNI 1 415

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.

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 jaw11 1 12 24 43.01 lower jaw 4 3 11 7 2 1 17 11 12.78 dm 3 max11 0 00 01 10.75 dm 3 mand000 000000.00 dm 4 max00 0 00 00 00.00 dm 4 mand110 000110.75 I1 max 00 0 00 00 00.00 I1 mand000 000000.00 I2 max 00 0 00 00 00.00 I2 mand111 100221.50 I3 max 11 4 41 16 64.51 I3 mand001 111221.50 C max 745.26 C mand 956.77 P1 max 00 0 00 00 00.00 P2 max 11 3 20 04 33.01 P3 max 00 3 20 03 22.26 P4 max 11 6 40 07 55.26 P2 mand320 000322.26 P3 mand112 100433.01 P4 mand003 200322.26 M1 mand323 211755.26 teeth ind. 96.77 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 21.50 axis 00.00 vertebrae 96.77 humerus 0 0 1 1 1 1 2 2 1.50 ulna 1 1 1 1 0 0 2 2 1.50 radius 1 1 1 1 0 0 2 2 1.50 metacarpus 2 21 1 10 03 22.26 metacarpus 3 00 0 00 00 00.00 metacarpus 4 11 1 10 02 21.50 metacarpus 5 11 3 20 04 33.01 femur 00.00 patella000 000000.00 tibia 21 1 10 03 22.26 fibula 00.00 astragalus 10.75 calcaneum 00.00 carpalia/tarsalia 43.01 metatarsus 2 11 1 10 02 21.50 metatarsus 3 00 0 00 00 00.00 metatarsus 4 00 1 10 01 10.75 metatarsus 5 11 1 10 02 21.50 phalanx 1 00 0 04 44 43.01 phalanx 2 00 0 01 11 10.75 phalanx 3 00 0 00 00 00.00 indeterminable 00.00

Total NISP 28 50 13 133 Total MNI 3 7 4 11

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.

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 jaw00 1 10 01 111.11 lower jaw00 1 10 01 111.11 dm 3 max00 0 00 00 00.00 dm 3 mand000 000000.00 dm 4 max00 0 00 00 00.00 dm 4 mand000 000000.00 I1 max 00 1 10 01 111.11 I1 mand000 000000.00 I2 max 00 0 00 00 00.00 I2 mand001 1001111.11 I3 max 00 0 00 00 00.00 I3 mand000 0001111.11 C max 1111.11 C mand 000.00 P1 max 00 0 00 00 00.00 P2 max 00 0 00 00 00.00 P3 max 00 0 00 00 00.00 P4 max 00 0 00 00 00.00 P2 mand000 000000.00 P3 mand110 0001111.11 P4 mand000 000000.00 M1 mand000 000000.00 teeth ind.000 000000.00 POSTCRANIAL NISP juv. MNIe juv. NISP subad. MNIe subad. NISP ad. MNIe ad. Total NISP Total MNIe % NISP atlas 00.00 axis 00.00 vertebrae 00.00 humerus 0 0 0 0 0 0 0 0 0.00 ulna 1 1 0 0 0 0 1 1 11.11 radius 1 1 0 0 0 0 1 1 11.11 metacarpus 2 00 0 00 00 00.00 metacarpus 3 00 0 00 00 00.00 metacarpus 4 00 0 00 00 00.00 metacarpus 5 00 0 00 00 00.00 femur 000.00 patella000 000000.00 tibia 00 0 00 00 00.00 fibula 00.00 astragalus 00.00 calcaneum 00.00 carpalia/tarsalia 00.00 metatarsus 2 00 0 00 00 00.00 metatarsus 3 00 0 00 00 00.00 metatarsus 4 00 0 00 00 00.00 metatarsus 5 00 0 00 00 00.00 phalanx 1 00 0 00 00 00.00 phalanx 2 00 0 00 00 00.00 phalanx 3 00 0 00 00 00.00 indeterminable 00.00

Total NISP 3 4 0 9 Total MNI1 102

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.

% NISP per skeletal element ‐ Goyet 80

NISP 40

% 20

Skeletal element A

% NISP per skeletal element ‐ Walsin 80

NISP 40

% 20

Skeletal element B

% NISP per skeletal element ‐ Hastière 80

NISP 40

% 20

Skeletal element C

Fig. 4.1: The percentage of NISP per skeletal elements per caves. A. Goyet, B. Walsin, C. Hastière.

% NISP per skeletal element without teeth ‐ Goyet 35 30 25 20 NISP 15 % 10 5 0

Skeletal element A

% NISP per skeletal element without teeth ‐ Walsin 35 30 25 20 NISP 15 % 10 5 0

Skeletal element B

% NISP per skeletal element without teeth ‐ Hastière 35 30 25 20 NISP 15 % 10 5 0

Skeletal element C

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

% NISP per skeletal element per horizon ‐ Goyet 80

NISP 40

% horizon 1 horizon 2 20 horizon 3 horizon 4

Skeletal element

% NISP per skeletal element per horizon ‐ Hastière 80

NISP 40

% horizon 1 horizon 2 20 horizon 5

Skeletal element

Fig. 4.3: The percentage NISP per skeletal elements per cave horizon: A. Goyet, B. Hastière

A C

% NISP without teeth per skeletal element ‐ Goyet, % NISP without teeth per skeletal element ‐ Goyet, horizon 1 horizon 3 20 20 15 15 NISP NISP 10 10

% % 5 5 0 0

Skeletal element Skeletal element

B D

% NISP without teeth per skeletal element ‐ Goyet, % NISP without teeth per skeletal element ‐ Goyet, horizon 2 horizon 4 20 20 15 15 NISP NISP 10 10

% % 5 5 0 0

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

% NISP per skeletal element without teeth‐‐Hastière, horizon 1 15

10 NISP

% 5

Skeletal element A

% NISP per skeletal element without teeth‐‐Hastière, horizon 2 15

10 NISP

% 5

Skeletal element B

% NISP per skeletal element without teeth‐‐Hastière, horizon 5 15

10 NISP

% 5

Skeletal element C

Fig. 4.5: The percentage NISP of Hastière per horizon: A. Horizon 1, B. Horizon 2, C. Horizon 5

4.2 Description of the fossil material

4.2.1 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) - 289.96 286 (2) - 253.96 136 (8) - 134.94 136 (12) - 111.52 125 (15) - 88.56 106 (25) - 52.61 57 (27) - 25.06 26 (28) - 21.50 24 (29) 68.86 69.22 - (34) - 118.12 97 (35) - 49.99 69 (36) - 71.93 75 (38) - 111.15 147 (39) - 92.15 134 (40) - 80.56 90

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).

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 4.4 – 4.7). 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 4.8-4.10). 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 2861-136-13

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 4.4-4.7). 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.

Number Site Horizon Plateau ID (1) (7) (9) (9a) (18) (19) (20) DP Goyet 3 157 2238 46 - 100.98 92.32 76.59 - - - Goyet 4 137 2842 83 ------31.03 Goyet 4 137 2842 80 - 99.92 88.96 73.29 - 64.7745.43 Goyet 4 137 2842 81 - 101.41 90.06 75.79 - 30.39 43.01 Goyet 4 137 2842 84 ------36.04 Goyet 4 136 2861 21 200.91 99.52 88.09 74.3 93.72 59.1 38.12 Goyet 4 137 2842 82 - 94.6 88.91 76.5 - 61.3440.13 Goyet 4 137 2842 93 - 97.5 87.9 72.58 - - 42.17 Goyet 4 137 2842 94 ------41.67 Walsin - 28 2480 3 - 92.72 86.01 72.86 - - 41.26 Walsin - 28 2480 8 - 103.87 91.88 76.31 - - 43.36 Walsin - 28 2480 4 ------38.34

Hastière 2 99 3872 59 - 92.88 90.1 78.52 - - 32.64 Hastière 2 99 3872 67 195.81 95.27 87.55 70.25 - 50.94 42.54 Hastière 2 99 3872 68 - - 90.19 77.57 - - 39.61 Hastière 2 99 3872 71 - - - - - 39.23 - Hastière 2 99 3872 78 - - - - - 43.03 -

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 2861- 136-21

Photo 4.4: Lower jaw with di2, di3, dc, dm2, dm3 and dm4 of a juvenile from Goyet (horizon 4) – specimen 2861-136-5a

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.

Number Position Crown Crown Cave Horizon Plateau ID Plateau Tooth DP in jaw length width Goyet 3 2238 2238 9 157 dm 3 upper left 21.65 8.12 Goyet 2 2829 2829 23 178 dm 4 - - - Goyet 3 2238 2238 15 157 dm 4 lower left 20.89 8.99 Hastière 2 3872 3872 58b 99 dm 3 upper left 21.05 9.28 Hastière 2 3872 3872 37 99 dm 4 lower left 20.14 8.51

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 Dental type n Mean Std dev

dm2 cl 5 8.33 1.48

dm2 cw 5 4.34 1.58 dm3 cl 2 22.17 0.74 dm3 cw 2 8.62 0.70 Goyet dm3 cl 4 11.05 3.17

dm3 cw 4 5.46 2.51

dm4 cl 3 16.51 3.87

dm4 cw 3 6.53 2.16

dm4 cl 2 19.96 0.25 Hastière dm4 cw 2 8.16 0.50

Table 4.14: All deciduous teeth from Goyet and Hastière.

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 4.1- 4.3). Table 4.15 gives the measurements of all incisors per cave.

Cave Dental type n Mean Std dev I1 cl 2 12.52 0.18 I1 cw 2 10.51 0.64 Goyet I3 cl 15 12.79 1.08 I3 cw 15 10.43 1.02 I1 cl 2 9.00 0.54 I1 cw 2 6.11 1.17 Walsin I3 cl 6 12.69 1.03 I3 cw 6 10.30 0.86

I2 cl 4 8.73 1.24

I2 cw 4 4.96 0.94 I3 cl 10 12.90 0.72 Hastière I3 cw 10 10.52 0.86

I3 cl 5 9.39 1.02

I3 cw 5 7.86 0.99

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 4.1- 4.3). 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.

Number Cave Number ID plateau position Crown length Crown width DP Walsin 2480 15 28 upper left 17.22 13.30 Walsin 2480 16 28 upper left 13.86 12.70 Walsin 2480 17 28 upper right 15.50 13.48 Walsin 2480 18 28 upper right 14.42 11.70 Walsin 2480 26 28 upper right 16.48 13.48 Walsin 2480 27 28 lower left 14.43 12.72 Walsin 2480 28 28 lower right 16.33 14.51 Walsin 2480 3 28 lower left mand 15.19 12.78 Walsin 2480 8 28 lower right mand 13.50 13.67

Table 4.16: The canines of Walsin and their measurements.

Cave Dental type n Mean Std dev 50 16.59 1.13 C max cl 50 13.38 0.93 C max cw Goyet 30 15.94 0.90 C mand cl 30 13.48 0.83 C mand cw 4 15.77 1.45 C max cl 4 13.24 0.37 C max cw Walsin 4 14.86 1.20 C mand cl 4 13.42 0.85 C mand cw 12 16.92 1.63 C max cl 12 13.69 0.89 C max cw Hastière 15 15.62 1.41 C mand cl 15 13.04 0.86 C mand cw

Table 4.17: The canines of the three caves.

Percentage of C per crown length class ‐ Goyet 50 40 (in%) 30

20 C max 10

Percentage C mand 0 13‐14 14‐15 15‐16 16‐17 17‐18 18‐19 19‐20 ? 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 30 %)

25 (in

20 15 10 C max 5 C mand Percentage 0 13‐14 14‐15 15‐16 16‐17 17‐18 18‐19 19‐20 ? 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 4.1- 4.3). 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. 4.8 - 4.9) 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.

Dental type n Mean Std dev P2 cl 3 16.93 0.94 P2 cw 3 12.85 1.12

P2 cl 27 16.68 0.90

P2 cw 27 12.28 1.04 P3 cl 24 24.48 1.35 P3 cw 24 17.96 1.00

P3 cl 41 22.63 0.86

P3 cw 41 16.28 0.84 P4 cl 11 40.02 1.54 P4 cw 12 21.56 1.47

P4 cl 38 24.02 0.97

P4 cw 38 14.99 0.73

Table 4.18: The premolars of Goyet.

Percentage of P3 per crown length class ‐ Goyet 50

%) 40

(in 30 P3 max 20 P3 mand 10 Percentage 0 21‐22 22‐23 23‐24 24‐25 25‐26 26‐27 27‐28 ? Crown length in classes (in mm)

Fig. 4.8: Frequency distribution of the crown length of the upper and lower P3’s.

Percentage of lower P4 per crown length class ‐ Goyet 50 %) 40 (in 30 20 10 Percentage 0 21‐22 22‐23 23‐24 24‐25 25‐26 Crown length in classes (in mm)

Fig. 4.9: Frequency distribution of the crown length of the lower P4’s.

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

2 P2 cl 9 16.42 0.67 P cl 8 16.67 1.11 2 P2 cw 9 12.37 1.09 P cw 8 12.26 1.14 3 P cl 5 23.55 1.04 P2 cl 19 16.50 0.60 3 P cw 5 17.42 0.85 P2 cw 19 12.29 0.61 3 P3 cl 11 22.83 1.17 P cl 15 23.34 1.63

3 P3 cw 11 16.72 1.14 P cw 15 17.11 0.99 4 P cl 3 42.56 1.18 P3 cl 24 22.40 1.12 4 P cw 3 20.83 0.24 P3 cw 23 16.45 1.03 4 P4 cl 12 23.24 1.18 P cl 12 41.70 2.26 4 P4 cw 12 14.73 0.73 P cw 12 21.41 1.27

P4 cl 20 23.66 1.08 Table 4.19: The premolars of Walsin. P4 cw 20 15.17 0.82

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 M1 cl 4 31.75 1.90 horizon 1 M1 cw 4 13.75 0.49 M1 cl 5 32.62 1.82 horizon 2 M1 cw 6 14.47 0.79 Goyet M1 cl 4 32.94 1.80 horizon 3 M1 cw 4 14.00 0.95 M1 cl 18 31.64 2.63 horizon 4 M1 cw 17 13.78 0.91 M1 cl 8 31.40 1.15 Walsin M1 cw 8 12.66 0.82 M1 cl 5 32.72 1.54 horizon 1 Hastière M1 cw 5 14.11 0.68

M1 cl 11 32.87 1.84 horizon 2 M1 cw 13 13.83 1.36

Table 4.21: The molars of all the caves.

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 30-31 class, the 33-34 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, 31-32 and 32-33. 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 33-34 class (fig. 4.10D). If we look at all the caves together, the highest number of teeth fall between the range of 30-34 mm.

A C

Percentage of M1 per cl class ‐ all caves Percentage of M1 per cl class ‐ Walsin 30 30 %) %)

25 25 (in (in 20 20

15 15 10 10 5 5 Percentage Percentage 0 0

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 30 30 %) %)

25 25 (in (in 20 20

15 15 10 10 5 5 Percentage Percentage 0 0

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.

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 4.22.

Site Horizon Number DP Plateau ID (Bd) (Kd) Goyet 3 2842 157 1 57.28 23.67 Goyet 3 2237 157 39 48.66 -

Table 4.22: Measurements of the humeri.

Photo 4.5: Humerus: left distal from Goyet (horizon 4) – specimen 2237-157-37

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 4.1- 4.3, 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 4 2861 136 27 88.71 8.68 11.42

Table 4.23: Measurements of the ulna.

D. RADIUS

In total eight radii were found (tables 4.1-4.3, fig. 4.1). Table 4.24 gives the available measurements of these radii.

Site Horizon Number DP Plateau ID (GL) (Bp) (Kd) (Bd) Goyet 2 2829 178 58 - - 21.2 - Goyet 4 2842 137 3 - - 23.2 - Walsin - 2480 28 72 221.36 31.21 23.66 45.57 Walsin - 2480 28 73 - 31.24 25.66 - Hastière 5 3895 54 A 1 - 36.2 - -

Table 4.24: Measurements of the radii.

Photo 4.6: Radius (left) with gnawing traces from Goyet (horizon 2) – specimen 2829-1872-58

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 4.1-4.3).

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 = 21.72 mm.

Site Horizon Number DP Plateau ID Element (GL) (Bp) (Kd) (Bd) Goyet 1 2812 190 18 MC2 - 13.54 - - Goyet 3 2237 157 24 MC2 80.6 14.54 12.38 14.15 Goyet 3 2237 157 27 MC2 81.28 15.96 13.12 15.5 Goyet 3 2237 157 25 MC2 75.3 14.7 12.53 13.79 Goyet 4 2861 136 37 MC2 78.58 14.42 12.32 15.62 Goyet 4 2861 136 38 MC2 76.03 - 11.18 14.54 Goyet 4 2861 136 39 MC2 - 11.01 10.11 - Hastière 2 3872 99 2i MC2 74.69 15.83 13.38 - Hastière 2 3872 99 3c MC2 78.86 15.82 11.97 15.72 Hastière 2 3872 99 1f MC2 - 14.83 - - Walsin - 2480 28 59 MC2 72.47 14.75 12.69 14.65 Walsin - 2480 28 60 MC2 80.7 15.6 12.92 15.39 Walsin - 2480 28 61 MC2 - 13.26 - - Goyet 1 2812 190 20 MC3 - 14.19 - - Goyet 3 2237 157 26 MC3 81.74 14.86 13.29 14.53 Goyet 4 2861 136 36 MC3 89.75 14.75 11.75 14.4 Goyet 3 2237 157 28 MC4 91.03 14.16 11.99 14.41 Goyet 3 2237 157 18 MC4 91.9 14.05 11.92 13.86 Goyet 3 2237 157 20 MC4 - 12.85 - - Goyet 4 2842 137 13 MC4 88.69 12.18 11.28 13.91 Goyet 4 2842 137 14 MC4 89.62 13.16 11.55 14.34 Goyet 4 2842 137 15 MC4 - - - 14.26 Hastière 2 3872 99 5c MC4 - - - 13.28 Hastière 2 3872 99 4b MC4 - 12.28 10.1 - Walsin - 2480 28 58 MC4 89.08 12.66 11.4 15.03 Goyet 3 2237 157 19 MC5 72.35 18.67 12.29 14.16 Goyet 4 2842 137 16 MC5 76.02 18.63 12.55 14.73 Goyet 4 2861 136 33 MC5 69.77 18.83 10.33 13.61 Goyet 4 2861 136 35 MC5 - - 10.83 13.61 Hastière 2 3872 99 6e MC5 64.64 13.45 12.21 10.78 Hastière 2 3872 99 7b MC5 75.47 17.54 11.66 13.95 Hastière 2 3872 99 8 MC5 71.66 16.59 13.09 15.4 Hastière 2 3872 99 9b MC5 68.27 16.13 12.79 15.61 Walsin - 2480 28 57 MC5 - 17.57 12.59 -

Table 4.25: Measurements of the metacarpals.

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 4 2842 137 5 190 47.17 18.81 39.71 Goyet 4 2842 137 6 - - - 40.39 Goyet 4 2861 136 30 - - 19.9 - Walsin - 2480 28 70 - - 18.33 41.36 Hastière 1 3863 95 21 - - - 11.56 Hastière 2 3872 99 20 - 42.19 16.87 -

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=16.32.

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.

Site Horizon Number DP Plateau ID Element (GL) (Bp) (Kd) (Bd) Goyet 3 2237 157 40 MT2 74.61 14.8 13.08 13.96 Goyet 4 2840 137 17 MT2 74.17 13.8 11.48 13.67 Hastière 2 3872 99 3d MT2 69.34 13.9 12.07 13.74 Goyet 3 2237 157 1 MT3 76.74 14.78 12.24 12.61 Goyet 3 2237 157 2 MT3 76.93 12.52 10.91 12.53 Goyet 3 2237 157 4 MT3 78.81 13.49 10.59 12.76 Goyet 3 2237 157 3 bis 29 MT3 83.13 15.27 11.76 13.03 Goyet 4 2840 137 18 MT3 76.06 12 11.03 11.39 Walsin - 2480 28 69 MT3 73.84 13.6 10.34 - Goyet 3 2237 157 35 MT4 78.17 11.49 10.16 12.08 Goyet 4 2861 136 31 MT4 58.3 11.28 8.26 9.04 Goyet 4 2840 137 19 MT4 81.98 10.76 10.12 12 Hastière 2 3872 99 1e MT4 74.49 10.39 10.79 10.88 Goyet 3 2237 157 30 MT5 63.4 13.3 8.77 12.88 Goyet 3 2237 157 34 MT5 - 10.62 - - Goyet 3 2237 157 36 MT5 82.62 17.97 8.7 11.57 Goyet 4 2861 136 32 MT5 54.26 10.67 7.94 9.51 Hastière 2 3872 99 4d MT5 - 13.51 10.42 - Hastière 2 3872 99 6d MT5 69.45 16.76 11.3 12.59 Walsin - 2480 28 68 MT5 63.5 13.04 8.32 11.16

Table 4.27: Measurements of the metatarsals.

4.3 Age determination

We made some frequency distributions on the wear-stages (STINER, 2004) of the fossil M1’s per cave (fig. 4. 11). 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 4.2.2.

Percentages of M1 per wear‐stages (Stiner) ‐ Goyet 30 %) 25 (in 20 15 10 5 Percentage 0 germ I II III IV V VI VII VIII IX ? Wear‐stages (according to stiner) A Percentages of M1 per wear‐stages (Stiner) ‐ Walsin 30

%) 25 20 (in

15 10 5 0 germ I II III IV V VI VII VIII IX ? Percentage Wear‐stages (according to stiner) B Percentages of M1 per wear‐stages (Stiner) ‐ Hastière 30

%) 25 20 (in

15 10 5 0 germ I II III IV V VI VII VIII IX ? Percentage Wear‐stages (according to Stiner) C

Fig. 4.11: The frequency distributions of the teeth per wear stages (Stiner). A. Goyet, B. Walsin, C. Hastière

Percentage of upper jaws per age class 120 %)

100 (in

80 60 Goyet 40 Walsin 20 Percentage 0 Hastière Juvenile Prime adult Old adult ? Age classes (according to stiner) A

Percentage of lower jaws per age class 100 %) 80 (in 60 Goyet 40 20 Walsin Percentage 0 Hastière juvenile prime adult old adult ? Age classes (according to Stiner) B

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.

Percentages of M1 per wear‐stages (Stiner) ‐ Recent material 30 %)

25 (in 20 15 10 5 Percentage 0 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.

4.4 Trace fossils

4.4.1 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

4.4.2 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).

Site Horizon Id Plateau Element Position Age Broken Weathering Gnawing traces Goyet 1 19 190 bone fragment - - x 1 proximal and distal Goyet 2 70 178 MT3 left juvenile x 1 proximal Goyet 2 69 178 MT4 left juvenile x 1 distal Goyet 2 58 178 radius left - - 1 proximal and distal Goyet 3 24 157 MC2 right adult - 1 proximal and distal Goyet 3 25 157 MC2 left sub-adult - 1 proximal and distal Goyet 3 21 157 MC5 right sub-adult x 1 proximal Goyet 3 5 157 humerus right adult x 1 distal Goyet 3 39 157 humerus right adult x 1 distal Goyet 3 8 157 ulna left juvenile x 1 proximal Goyet 3 11 157 ulna left adult x 1 proximal Goyet 4 39 136 MC2 right juvenile - 1 proximal and distal Goyet 4 35 136 MC5 right sub-adult - 1 proximal en diaphyse Goyet 4 32 136 MC5 right sub-adult - 1 proximal and distal Goyet 4 3 137 radius right - - 1 proximal and distal Walsin 57 28 MC5 left juvenile - 1 distal Walsin 72 28 radius right adult - 1 diaphyse Hastière 1 19 95 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.

Percentage skeletal elements with gnaw‐traces ‐ per cave and per horizon 8

%) 7 6 (in 5 4 3 2 1 0 Percentage

Skeletal elements with gnawing traces: percentages per age class 40 %) 30 (in

10 Percentage 0 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 2842-137-5

4.6 Inventory of the recent material

For the recent material we measured 33 specimens. Table 4.30 gives an overview of these specimens. Twenty-two specimens were collected in DR Congo, one in Rwanda and one in South Africa. Seven specimens come from unknown locations of which two lived and/or died in captivity. There were also two hyenas that lived in the zoo of Sint-Agatha-Berchem.

Location ID Year Sex Age Element skull Congo, Katanga 9967 1941-1943 unknown old adult lower jaw skull Congo, Katanga 11799 20/10/1947 female old adult lower jaw skull Congo, Katanga 11800 20/04/1948 male prime adult lower jaw skull Congo, Katanga 11801 20/04/1948 male old adult lower jaw skull Congo, Katanga 11802 14/04/1948 female juvenile lower jaw skull Congo, Katanga 11803 3/12/1947 male juvenile lower jaw skull Congo, Katanga 11804 1941-1943 unknown prime adult lower jaw skull Congo, Kivu 4609 28/02/1938 male juvenile lower jaw skull Congo, Kivu 4610 28/02/1938 male juvenile lower jaw skull Congo, Kivu 4611 4/03/1938 male prime adult lower jaw skull Congo, Kivu 4612 22/04/1938 female old adult lower jaw skull Congo, Kivu 8632 unknown unknown prime adult lower jaw skull Congo, Kivu 8633 unknown unknown prime adult lower jaw skull Congo, Kivu 8635 12/03/1952 unknown prime adult lower jaw skull Congo, Kivu 9480 28/01/1949 unknown prime adult lower jaw skull Congo, Kivu 10250 3/06/1948 unknown prime adult lower jaw skull Congo, Kivu 21436 6/05/1954 male old adult lower jaw skull Congo, lake Albert 10336 feb/53 male prime adult lower jaw

skull Congo, Masuku 35152 unknown unknown prime adult lower jaw skull Congo, Ubangi Liki 7705 14/04/1948 male prime adult lower jaw skull Congo, Uélé 21278 6/05/1954 female old adult lower jaw skull Congo, Uélé 21302 11/06/1952 female prime adult lower jaw skull Rwanda, Kagera 8634 feb/47 unknown prime adult lower jaw skull South Africa 1007-B det. 1935 unknown old adult lower jaw skull unknown 7602 14/08/1951 male prime adult lower jaw skull unknown 12662 feb/53 female prime adult lower jaw skull unknown 22476 24/10/1995 unknown prime adult lower jaw skull unknown 1008-C unknown unknown prime adult lower jaw skull unknown 1008-E male 1897 old adult lower jaw skull unknown - captivity 3336 det. 1956 male prime adult lower jaw skull unknown - died in captivity 7601 14/08/1951 male prime adult lower jaw skull zoo of St-Agatha Berchem 7546 10/05/1940 unknown prime adult lower jaw skull zoo of St-Agatha Berchem 7547 10/05/1940 unknown old adult lower jaw

Table 4.30: The inventory of the recent material.

4.7 Description of the recent material

4.4.1 Cranial

A. CRANIUM and MAXILLA

In total there are 33 skulls and most individuals belong to the prime adult-age class (fig. 4.15). Table 4.31 gives all available measurements.

Percentage of the skulls per age class ‐ recent material 100 %) 80 (in 60 40 20 Percentage 0 juvenile prime adult old adult Age classes (according to Stiner)

Fig. 4.15: The percentages of the recent skulls per age class.

Location ID (1) (2) (8) (12) (15) (25) (27) (28) (29) (34) (35) (36) (38) (39) (40) Congo, Katanga 9967 292.73 ‐ 109.72 94.17 85.88 ‐‐‐ 73.45 82.28 68.26 68.17 ‐‐‐ Congo, Katanga 11804 ‐‐ 106.36 80.91 82.06 ‐‐‐‐ 77.46 63.63 62.39 ‐‐‐ Congo, Katanga 11803 222.34 191.86 85.06 83.40 92.72 55.61 26 .27 18.12 72.50 82,08* 49,41* 48,24* 90.38 87.51 66.96 Congo, Katanga 11802 235.99 207.47 89.88 85.86 85.65 53.04 21.42 19.63 71.92 69.26 55.54 57.27 96.93 91.90 71.77 Congo, Katanga 11799 277.39 226.24 104.27 81.76 80.38 52.93 25.01 19.39 70.95 80.32 60.59 66.43 121.59 98.83 76.06 Congo, Katanga 11801 288.68 237.80 108.62 93.96 87.01 53.64 24.91 22.11 72.80 8 2.43 65.50 68.16 119.05 110.83 86.43 Congo, Katanga 11800 223.77 193.09 83.28 76.97 91.02 51.38 27.93 19.97 77.64 71.26 50.21 50.50 88.53 89.05 88.06 Congo, Kivu 10250 242.97 201.56 94.06 78.67 70.39 4 8.86 24.78 22.21 74.45 68.60 54.52 54.03 ‐‐ 73.96 Congo, Kivu 8635 269.93 ‐ 93.34 84.52 82.28 ‐‐‐ 72.12 ‐‐‐‐‐‐ Congo, Kivu 9480 218.51 231.63 97.87 77.54 80.13 52.47 24.81 23.01 67.01 74. 35 61.15 61.46 126.70 105.98 83.85 Congo, Kivu 8632 269.19 227.95 97.95 74.11 80.82 52.80 25.89 25.13 76.54 75. 35 57.55 58.13 116.36 101.18 72.83 Congo, Kivu 8633 256.12 218.23 87.90 70.06 78.68 52.58 21.90 70.89 71.52 54.97 56.04 116.75 101.15 77.53 Congo, Kivu 4611 197.16 173.13 69.26 72.25 ‐ 46.57 26.90 18.64 68.24 66.95 45.58 46.37 80.92 78.05 62.94 Congo, Kivu 4612 231.28 229.10 102.58 89. 40 83.50 50.47 25.67 22.97 68.44 75.49 59.34 60.67 ? 119.75 103.74 Congo, Kivu 4609 85.13 ‐ 28.88 26.21 ‐‐‐‐ 43.02 32.03 23.50 24.24 43.54 ‐‐ Congo, Kivu 4610 ‐‐ 28.29 23.34 ‐‐‐‐ 44.76 33.60 24.23 25.17 ‐‐‐ Congo, Kivu 21436 272.03 22.73 100.48 93.23 80.40 51.99 26.4 7 21.25 73.61 83.27 58.04 59.15 116.11 99.67 82.38 Congo, lake Albert 10336 277.68 229.63 ‐‐ 82.75 48.32 24.02 19.48 77.85 77. 34 62.51 64.32 120.26 104.22 81.23 Congo, Ubangi 7705 270.44 ‐ 97.09 84.46 79.23 ‐‐‐ 80.57 78.09 62.96 63.23 ‐‐‐ Congo, Uélé 21302 268.43 224.05 100.05 77.00 78.26 53.24 25.3 6 19.01 72.29 75.18 56.88 60.27 122.22 101.84 80.96 Congo, Uélé 21278 297.63 244.68 110.12 101.87 83.30 53.95 26 .15 19.59 76.16 85.39 63.39 67.77 133.25 108.97 79.81 Congo,Masuku 35152 274.25 228.71 99.12 84.30 82.77 53.49 26.17 20.08 71.55 78.20 58.86 61.49 124.41 105.89 84.27 Rwanda, Kagera 8634 260.35 216.79 95.85 76.92 80.53 53.70 27 .44 21.76 75.49 73.78 53.97 57.43 111.75 93.48 72.14 South ‐ Africa 1007 ‐ B 295.27 ‐ 102.60 88.49 86.89 54.52 24.43 27.02 82 .11 81.12 64.28 66.72 114.69 84.11 77.18 unknown 22476 244.19 212.13 100.35 74.26 76.08 46.29 22 .07 18.32 75.00 70.64 55.40 58.71 104.07 88.14 76.33 unknown 12662 268.84 224.29 103.38 95.69 82.13 54.93 23 .16 17.15 78.11 84.59 62.70 63.98 122.46 100.33 77.77 unknown 7602 221.01 ‐ 86.58 73.43 71.77 45.70 22.76 18.31 71 .06 65.38 50.41 54.38 94.22 83.72 66.76 unknown 1008 ‐ C 263.10 229.50 93.49 82.36 79.61 50.45 23.87 1 8.07 67.03 96.16 58.96 60.98 114.25 94.74 71.88 unknown 1008 ‐ E 269.54 230.52 103.28 91.14 81.19 49.19 25.29 19. 39 65.56 90.89 56.39 58.66 114.78 100.28 78.13 unknown ‐ captivity 3336 251.56 214.87 98.34 86.64 79.33 49.92 22 .87 20.67 68.70 74.64 56.66 57.24 116.06 100.87 75.42 unknown ‐ died in captivity 7601 242.17 207.30 93.42 87.38 78.86 51.46 26.36 20.06 65.23 72.15 52.02 55.46 103.46 96.13 78.19 zoo of St ‐ Agatha Berchem 7546 245.09 212.00 95.97 69.11 ‐ 50.17 24.26 24.14 57.87 55.18 51.90 52.43 105.39 83.45 54.68 zoo of St ‐ Agatha Berchem 7547 ‐‐ 98.11 85.37 82.05 ‐‐‐‐ 88.11 60.39 63.13 ‐‐‐

Table 4.31: All the skull measurements of the recent material.

B. MANDIBULA

Table 4.32 gives a complete overview of the lower jaws and the available measurements.

Location ID (1) (7) (9) (9a) (18) (19) (20) Congo, Katanga 9967 194.73 96.87 85.83 74.28 90.16 54.46 38.47 Congo, Katanga 11799 176.37 93.14 83.26 72.62 82.94 47.80 34.84 Congo, Katanga 11800 150.95 87.22 76.95 64.19 65.72 38.78 30.70 Congo, Katanga 11801 187.44 97.65 87.65 76.03 86.53 51.3 38.36 Congo, Katanga 11802 162.64 90.31 85.44 76.24 70.50 41.48 29.76 Congo, Katanga 11803 148.54 83.87* 75.91* - 66.24 38.74 - Congo, Katanga 11804 181.63 92.37 82.26 69.09 90.91 53.65 37.21 Congo, Kivu 4609 52.82 - - - 23.46 - - Congo, Kivu 4610 55.56 - - - 22.77 - - Congo, Kivu 4611 139.16 79.89 69.93* 57.11* 59.89 28.95 28.60* Congo, Kivu 4612 ------Congo, Kivu 8632 177.57 901.11 80.29 65.22 84.13 49.23 31.54 Congo, Kivu 8633 165.25 87.61 77.9 64.34 83.37 48.54 29.92 Congo, Kivu 8635 179.50 92.36 82.67 66.56 80.39 49.61 33.43 Congo, Kivu 9480 186.41 92.37 83.24 67.03 83.59 48.17 34.33 Congo, Kivu 10250 161.58 82.11 73.23 61.64 75.06 41.95 28.43 Congo, Kivu 21436 177.55 91.17 80.93 67.25 79.91 47.65 33.71 Congo, lake Albert 10336 118.12 90.37 79.84 66.62 82.24 51.14 34.05 Congo, Ubangi 7705 181.09 89.32 79.82 68.79 89.82 52.12 31.13 Congo, Uélé 21278 193.07 96.80 87.47 77.37 91.42 58.43 39.76 Congo, Uélé 21302 171.58 88.57 79.60 64.12 89.83 52.50 31.88 Congo, Masuku 35152 176.76 93.33 80.75 66.64 85.57 51.47 33.43 Rwanda, Kagera 8634 177.50 89.92 81.38 64.48 81.55 48.12 31.49 South-Africa 1007-B 183.58 93.22 81.62 68.96 86.99 50.69 37.99 unknown 7602 152.51 80.62 74.25 63.40 63.26 33.08 25.74 unknown 12662 175.97 90.67 80.30 70.35 80.13 53.07 38.00 unknown 1008-C 177.49 88.48 79.02 67.73 79.88 48.19 37.74 unknown 1008-E 182.07 90.51 80.28 68.11 79.36 49.56 36.53 unknown - died in captivity 7601 167.06 87.47 80.01 69.31 71.49 40.30 29.62 unkown 22476 167.88 85.16 78.38 65.53 77.19 50.95 30.91 unkown - captivity 3336 169.91 86.63 77.64 65.95 73.66 48.83 33.53 Zoo of St-Agatha Berchem 7546 ------zoo of St-Agatha Berchem 7547 175.28 90.82 81.75 70.73 80.21 49.67 36.31

Table 4.32: All the available measurements of the recent lower jaws. * In this case, measurements for resp. dm2 and dm3 are given instead of the P2 and P3.

Tables 4.33 and 4.34 give the available measurements of the lower jaws per sex; 19 individuals have a known gender: six females and 13 males.

Location ID Sex (1) (7) (9) (9a) (18) (19) (20) Congo, Katanga 11799 F 176.37 93.14 83.26 72.62 82.94 47.80 34.84 Congo, Katanga 11802 F 162.64 90.31 85.44 76.24 70.50 41.48 29.76 Congo, Kivu 4612 F ------Congo, Uélé 21278 F 193.07 96.80 87.47 77.37 91.42 58.43 39.76 Congo, Uélé 21302 F 171.58 88.57 79.60 64.12 89.83 52.50 31.88 unknown 12662 F 175.97 90.67 80.30 70.35 80.13 53.07 38.00

Table 4.33: The available measurements of the recent lower female jaws.

Location ID Sex (1) (7) (9) (9a) (18) (19) (20) Congo, Katanga 11800 M 150.95 87.22 76.95 64.19 65.72 38.78 30.70 Congo, Katanga 11801 M 187.44 97.65 87.65 76.03 86.53 51.3 38.36 Congo, Katanga 11803 M 148.54 83.87* 75.91* - 66.24 38.74 - Congo, Kivu 4609 M 52.82 - - - 23.46 - - Congo, Kivu 4610 M 55.56 - - - 22.77 - - Congo, Kivu 4611 M 139.16 79.89 69.93* 57.11* 59.89 28.95 28.60* Congo, Kivu 21436 M 177.55 91.17 80.93 67.25 79.91 47.65 33.71 Congo, lake Albert 10336 M 118.12 90.37 79.84 66.62 82.24 51.14 34.05 Congo, Ubangi 7705 M 181.09 89.32 79.82 68.79 89.82 52.12 31.13 unknown 7602 M 152.51 80.62 74.25 63.40 63.26 33.08 25.74 unknown 1008-E M 182.0790.51 80.28 68.11 79.36 49.56 36.53 unknown - died in captivity 7601 M 167.06 87.47 80.01 69.31 71.49 40.30 29.62 unkown - captivity 3336 M 169.91 86.63 77.64 65.95 73.66 48.83 33.53

Table 4.34: The available measurements of the recent lower male jaws. * In this case, measurements for resp. dm2 and dm3 are given instead of the P2 and P3.

C. DENTES

Deciduous teeth

Quite a number of deciduous teeth were measured. Not only deciduous molars were present, but also deciduous incisors and canines (not found among the fossil material). Table 4.35 gives an overview of the measured deciduous teeth and the calculated means and standard deviations.

Dental type n Mean Std dev di1 cl 2 2.71 0.16 di1 cw 2 1.81 0.13

di1 cl 2 1.71 0.23

di1 cw 2 1.20 0.23 di2 cl 2 2.90 0.01 di2 cw 2 2.77 0.07

di2 cl 2 2.30 0.08

di2 cw 2 1.84 0.26 di3 cl 2 2.98 0.64 di3 cw 2 3.32 0.47

di3 cl 2 2.56 0.07

di3 cw 2 2.95 0.37 dC max cl 2 3.59 0.81 dC max cw 2 2.95 1.15 dC mand cl 4 5.28 1.52 dC mand cw 4 3.42 0.45 dm2 cl 3 10.64 0.81 dm2 cw 3 5.80 0.49

dm2 cl 3 9.15 0.54

dm2 cw 3 4.96 0.68

dm3 cl 3 15.39 2.79

dm3 cw 3 5.52 0.46

Table 4.35: The deciduous teeth of the recent material.

Incisors

Table 4.36 gives the calculated means and standard deviations of the measured incisors.

Dental type n Mean Std dev I1 cl 28 6.72 0.57 I1 cw 28 4.64 0.45

I1 cl 27 4.99 0.63

I1 cw 27 3.04 0.50 I2 cl 27 8.43 0.74 I2 cw 27 5.83 0.58

I2 cl 25 6.71 0.79

I2 cw 26 4.32 0.47 I3 cl 26 12.04 1.12 I3 cw 26 9.06 1.34

I3 cl 27 7.90 0.80

I3 cw 27 7.34 0.91

Table 4.36: The incisors of the recent material.

Canines

The calculated means and standard deviations of the canine measurements can be found in table 4.37. Tables 4.38 and 4.39 give the measurements per sex.

Dental type n Mean Std dev C max cl 26 14.58 1.51 C max cw 26 11.62 0.84 C mand cl 26 14.83 1.22 C mand cw 26 11.05 0.81 Table 4.37: The canines of the recent material.

Dental type n Mean Std dev Dental type n Mean Std dev C max cl 4 14.67 1.68 C max cl 8 14.36 1.26 FEMALE C max cw 4 11.47 0.91 MALE C max cw 8 11.03 0.94 C mand cl 6 15.49 2.15 C mand cl 8 14.18 0.89 C mand cw 6 11.66 0.94 C mand cw 8 11.62 0.71

Table 4.38: The female canines of the recent Table 4.39: The male canines of the recent material. material.

For the fossil material, we made graphs (fig. 4.6 and 4.7) displaying the percentages of the upper and lower canines found in Goyet and Hastière per crown length classes and per crown width classes. For these graphs we included the teeth still in the jaws and the isolated canines. Such graphs for the recent material are shown in fig. 4.16. Looking at this graph we see that most upper and lower canines have a crown length between 14 and 15 mm (fig. 4.16A). For the crown width we see that most upper canines have a width between 10 and 11 mm, but the lower canines are somewhat bigger with most canines having a width between 11 and 12 mm (fig 4.16B).

Figure 4.17 shows the percentages of crown length and crown width per classes and per sex. We see no clear difference between males and females.

Percentage of C per crown length class ‐ recent material 30 25 (in%) 20 15 10 C mand 5

Percentage C max 0 12:13 13‐14 14‐15 15‐16 16‐17 17‐18 18‐19 19‐20 ? crown length per classes (in mm) A

Percentage of C per crown width class, recent material 40

30 (in%)

20 C mand 10

Percentage C max 0 9‐10 10‐11 11‐12 12‐13 13‐14 ? crown width per classes (in mm) B

Fig. 4.16: A. The percentages of the crown length of C of the recent material per crown length classes. B. The percentages of the crown width C of the recent material per crown length class.

A C

Percentage of upper C per crown length class per sex ‐ Percentage of lower C per crown length class per sex ‐ recent material recent material 40 40 %) %)

30 30 (in (in

20 20 Female Female 10 10 Male Male percentage percentage 0 0 12‐13 13‐14 14‐15 15‐16 16‐17 17‐18 ? 12‐13 13‐14 14‐15 15‐16 16‐17 17‐18 18‐19 19‐20 crown length per classes (in mm) crown length per classes (in mm)

B D

Percentage of upper C per crown width class per sex ‐ recent Percentage of lower C per crown width class per sex ‐ recent material material 40 60 %) %)

50 30 (in (in

40 20 30 Female 20 Female 10 10 Male Male percentage 0 percentage 0 9‐10 10‐11 11‐12 12‐13 ? 10‐11 11‐12 12‐13 13‐14 crown length per classes (in mm) crown length per classes (in mm)

Fig.4.17: The percentages of C of the recent material per crown length and width classes per sex. A. Upper crown length B. Upper crown width C. Lower crown length D. Lower crown width.

Premolars

Also for the premolars we calculated the means and standard deviations (tables 4.40-4.42).

Dental type n Mean Std dev p1 cl 28 7.33 0.72 p1 cw 28 6.64 0.67 p2 cl 28 14.83 1.10 p2 cw 28 10.68 1.23

p2 cl 26 14.55 0.90

p2 cw 26 9.49 1.91 p3 cl 27 21.40 1.41 p3 cw 27 15.89 1.07

p3 cl 26 20.03 1.14

p3 cw 27 13.82 1.00 p4 cl 29 35.04 2.19 p4 cw 29 19.58 1.38

p4 cl 27 21.61 1.45

p4 cw 28 12.30 0.98

Table 4.40: The premolars of the recent material .

Dental type n Mean Std dev Dental type n Mean Std dev p1 cl 4 7.84 1.28 p1 cl 11 7.21 0.69 p1 cw 4 7.23 1.14 p1 cw 11 6.52 0.57 p2 cl 6 15.48 0.73 p2 cl 9 14.16 1.31

2 p2 cw 6 11.35 0.99 p cw 9 9.97 1.67

p cl 5 14.94 0.67 p2 cl 8 14.66 1.03 2 p cw 5 10.53 0.65 p2 cw 8 9.66 0.85 2 3 Female p3 cl 6 22.12 1.33 Male p cl 8 20.80 1.73 3 p3 cw 6 16.74 1.07 p cw 8 15.51 1.20

p cl 5 20.18 1.01 p3 cl 8 18.22 1.46 3 p cw 6 14.58 0.83 p3 cw 8 13.04 1.03 3 4 p4 cl 6 36.03 2.03 p cl 10 35.44 2.24

4 p4 cw 6 20.48 0.78 p cw 9 19.38 1.57

p cl 6 22.46 1.91 p4 cl 7 19.77 1.65 4 p cw 6 13.01 1.07 p4 cw 7 11.48 1.18 4

Table 4.41: The female premolars of the recent Table 4.42: The male premolars of the recent material. material.

For the fossil material we made some graphs (Goyet) that display the percentages of P3 per crown length and crown width classes. We took the P3 because it was the most numerous tooth. The graphs for the recent material are shown in fig. 4.18 and show the different percentages of the crown length classes. In these graphs we can see that both the upper and lower P3’s show unimodal curves. The lower jaw teeth are somewhat smaller than the upper-jaw teeth: the upper-jaw teeth have a higher crown length compared to the lower jaw teeth. But we have to keep in mind here that the female upper teeth have the highest percentage at the 22-23 mm class and that this has an effect on the percentage of all teeth combined.

Percentage of upper P3 per crown length class ‐ recent material

60 %) 50 (in 40 30 Unknown 20 Female 10 Percentage 0 Male 17‐18 18‐19 19‐20 20‐21 21‐22 22‐23 23‐24 24‐25 ? crown lenght per classes (in mm) A

Percentage of lower P3 per crown length class ‐ recent material 60 %)

50 (in 40 30 unknown 20 Female 10 Percentage 0 Male 17‐18 18‐19 19‐20 20‐21 21‐22 22‐23 ? crown lenght per classes (in mm) B

Fig. 4.18: A. Frequency distributions of the crown length of the recent upper P3’s of the male, female skulls and skulls with unknown gender, B. Frequency distributions of the crown length of the recent lower P3’s of the male, female skulls and skulls with unknown gender.

Molar

Table 4.43 presents the means and standard deviations of the recent M1’s measured. Tables 4.44 and 4.45 give these calculations for the two sexes. The graph of the crown lengths per classes is shown in fig. 4.19.

Dental type n Mean Std dev M1 cl 29 27.34 1.80 M1 cw 29 11.22 0.84

Table 4.43: The M1’s of the recent material.

Dental type n Mean Std dev Dental type n Mean Std dev Female M1 cl 6 28.56 2.55 Male M1 cl 9 27.70 1.56 M1 cw 6 11.71 1.35 M1 cw 9 11.34 0.57

Table 4.44: The female M1’s of the recent material. Table 4.45: The male M1’s of the recent material.

Most molars have a crown width between 27 and 28 mm, and a crown width between 11 and 12 mm (fig. 4.19).

Percentage of M1 per crown length class ‐ recent material 40 %) 30 (in

20 Unknown 10 Female Percentage 0 Male 24‐25 25‐26 26‐27 27‐28 28‐29 29‐30 30‐31 31‐32 ? crown length per classes (in mm) A

Fig. 4.19: Frequency distributions of the crown length of M1’s of the male, female skulls and skulls with unknown gender.

4.8 The fossil and recent spotted hyena

The complete results of the performed ANOVA’s are given in annex. The significance level was set on p < 0.01 for all statistical tests. For the fossil material we compared the tooth- measurements of the three caves to see if there were significant differences. The F-values and P-values of these tests are given in tables 4.53 and 4.54 and the results show that we can group the crown lengths and crown widths of the fossil teeth together except for the crown width of M1 as it has a p-value of 0.0001.

Tooth Measurement F-value P-value Tooth Measurement F-value P-value

P3 crown length 3.19 0.0513 P2 crown length 0.49 0.6163

P3 crown width 3.58 0.0369 P2 crown width 0.03 0.9671

P4 crown length 3.26 0.0567 P3 crown length 0.80 0.4519

P4 crown width 0.35 0.7086 P3 crown width 0.93 0.4000

P4 crown length 2.82 0.0664 Table 4.53: The F-values and P-values of the ANOVA’s on the fossil upper teeth. P4 crown width 1.30 0.2792

M1 crown length 1.86 0.1659

M1 crown width 10.67 0.0001

Table 4.54: The F-values and P-values of the ANOVA’s on the fossil lower teeth.

For the recent material we first distinguished between wild and captive spotted hyenas as there may be differences among their development (BINDER & VALKENBURGH, 2000; VAN HORN et al., 2005). All wild spotted hyenas are put into a ‘W-group’, the captive animals are put together in a ‘C-group’. Our W-group consists of animals form different locations and countries, therefore we performed ANOVA’s to see if there were significant differences among these animals. Table 4.56 shows the different groups; the results of the ANOVA’s are shown in tables 4.57-4.59. The results of the ANOVA’s show that there are no significant differences between the wild hyenas, so they can be indeed grouped together as one W-group. As there are only four captive hyenas, we did not perform an ANOVA on this group. Next, we performed ANOVA’s to see if there were significant differences between the two recent groups (tables 4.60 and 4.61); we found no significant differences between the wild and captive hyenas.

Location Group Congo, Katanga 1 Congo, Kivu 2 Congo, lake Albert 3 Congo, Ubangi 4 Congo, Uélé 5 Congo,Masuku 6 Rwanda, National Park 7 South Africa 8 unknown 9 Table 4.56: Wild recent hyenas groups per location and country.

Measurement F-value P-value W- Total skull group 0.71 0.6811 length

Table 4.57: The F-value and P-value of the ANOVA on the recent skull material.

Tooth Measurement F-value P-value Tooth Measurement F-value P-value

P2 crown length 0.90 0.4205 P2 crown length 1.73 0.1957

P2 crown width 0.11 0.8953 P2 crown width 0.27 0.8441

P3 crown length 2.60 0.0995 P3 crown length 2.23 0.1192

P3 crown width 0.73 0.4962 P3 crown width 1.30 0.3036

P4 crown length 1.18 0.3270 P4 crown length 0.43 0.7308

P4 crown width 1.24 0.3107 P4 crown width 1.23 0.3269

M1 crown length 1.05 0.3695 Table 4.58: The F-values and P-values of the ANOVA’s on the recent upper teeth of the W- M1 crown width 2.42 0.1001 group.

Table 4.59: The F-values and P-values of the ANOVA’s on the recent lower teeth of the W- group.

Tooth Measurement F-value P-value Tooth Measurement F-value P-value

P2 crown length 0.17 0.6813 P2 crown length 2.98 0.0970

P2 crown width 0.23 0.6333 P2 crown width 0.00 0.9744

P3 crown length 1.09 0.3059 P3 crown length 2.55 0.1237

P3 crown width 0.10 0.7573 P3 crown width 0.09 0.7696

P4 crown length 0.00 0.9976 P4 crown length 0.09 0.7721

P4 crown width 0.69 0.4157 P4 crown width 1.23 0.2783

M1 crown length 0.12 0.7349 Table 4.60: The F-values and P-values of the ANOVA’s on the recent upper teeth. M1 crown width 0.09 0.7691

Table 4.61: The F-values and P-values of the ANOVA’s on the recent lower teeth.

Tables 4.62 and 4.63 give the results of the ANOVA’s and Post hoc tests performed to compare the fossil and recent teeth measurements. Although we found no significant differences between the two recent groups, we kept both groups separately. This gives a somewhat surprising result: While both recent groups do not differ significantly from each other, the W- group differs significantly from the fossil group, but this is not straightforward for the C-group. For this latter we find tooth measurements that differ significantly and others that do not (tables 4.62- 4.63).

ANOVA Tukey-Kramer Post hoc Multiple Comparison test Teeth Fossil-Wild-Captive P-value F-value P-value F-W F-C W-C P2 cl 12.14 <0.0001 <0.0001 0.0308 0.9055 P2 cw 8.29 0.0011 0.0007 0.1107 0.8734 P3 cl 26.43 <0.0001 <0.0001 0.0002 0.5758 P3 cw 21.95 <0.0001 <0.0001 0.0174 0.9443 P4 cl 34.57 <0.0001 <0.0001 0.0015 0.9993 P4 cw 12.35 <0.0001 <0.0001 0.3246 0.6711 Table 4.62: The F-values and P-values of the ANOVA’s on the fossil and recent upper individual and upper- jaw teeth.

ANOVA Tukey-Kramer Post hoc Multiple Comparison test Teeth Fossil-Wild-Captive P-value F-value P-value F-W F-C W-C P2 cl 59.31 <0.0001 <0.0001 0.0025 0.2220 P2 cw 38.88 <0.0001 <0.0001 0.0013 0.9031 P3 cl 64.38 <0.0001 <0.0001 <0.0001 0.8687 P3 cw 68.57 <0.0001 <0.0001 <0.0001 0.8404 P4 cl 33.04 <0.0001 <0.0001 0.1806 0.9281 P4 cw 100.27 <0.0001 <0.0001 0.3246 0.9650 M1 cl 53.75 <0.0001 <0.0001 <0.0001 0.9471 Table 4.63: The F-values and P-values of the ANOVA’s on the fossil and recent lower individual and lower-jaw teeth.

Kurten (1965) used the crown length of the lower P2 and lower P3 to distinguish between Eemian and Weichselian hyenas. We made scatter plots of the P2’s and P3’s of the lower jaws accordingly to compare the fossil and recent hyena (fig. 4.20). A scatter plot of the crown length and crown width of the lower P3 was also made (fig 4.21). Finally we plotted the total skull length against the crown length of P4, the result is shown in fig. 4.22.

17 P2

16 lower

15 Fossil length 14 W‐group C‐group Crown 13

12 17 18 19 20 21 22 23 24 25 Crown length lower P3

Fig. 4.20: scatter plot of the crown length of lower P2’s and lower P3’s.

17 P3

16 lower 15 Fossil 14 length W‐group 13 C‐group Crown 12 11 17 18 19 20 21 22 23 24 25 Crown length lower P3

Fig. 4.21: scatter plot of the crown length and crown width of the P3’s

38 P4

33 lenght

Fossil 28 W‐group crown 23 C‐group

18 210 230 250 270 290 310 Total skull length

Fig. 4.22: Scatter plot of the total skull length against the crown length of P4.

5. DISCUSSION

5.1 Taphonomy

The assemblages from Goyet, Walsin and Hastière all show a clear dominance of isolated teeth (fig. 4.1). This can be explained by the fact that teeth consist of hard material (enamel and dentin) which leads to a better preservation. The second most preserved element found in the three caves is the lower jaw. Upper jaws, carpals, metacarpals, vertebrae, metatarsals, tarsals and phalanges are skeletal elements also well represented in the three caves (fig. 4.1-4.2). The individual teeth and lower jaws of the hyena material of the Spy cave (Namur province, Belgium) are also the best represented skeletal elements (GERMONPRÉ et al., in press).

Fig. 5.1: The % NISP per skeletal elements of the Spy cave. (GERMONPRÉ et al., in press)

First, we will discuss the hyena taphonomy per cave and per cave horizon; then we will discuss the general taphonomy.

GOYET

Horizon 1 has a total NISP of 43 hyenas, which is not so numerous. Besides individual teeth, only four different skeletal elements (lower jaws, carpals/tarsals, metacarpals, and phalanges) and quite some equivocal elements were found (fig. 4.4A). Our calculated MNI is five, this is also the MNI calculated by Dupont (GERMONPRÉ, 1997), so it would seem that few or no material was lost. Dupont (1906) described the hyena assemblage in this horizon as ‘un court séjour’ because he found only six bones with gnawing traces of hyenas. The hyena assemblage from horizon 2 has a total NISP of 71 and, apart from the teeth, seven different identified skeletal elements were found: upper jaws, lower jaws, vertebrae, radii, carpals/tarsals, metatarsals and phalanges. Also here, quite some indeterminable elements were found (fig. 4.4B). The card of Dupont with information on the bones found in this horizon is missing.

Dupont (1906) identified 14 individuals in horizon 3 and this corresponds with our calculated MNI (table 4.6). He further mentions that a great number of bones indicate that the cave was longer inhabited by hyenas. Our results point in the same direction. First, we have a lot more bones than in horizon 1 or horizon 2. Second, the hyena assemblage is also more diverse: we found thirteen different skeletal elements (fig. 4.4C). Dupont (1906) states that some of these hyenas were probably killed by cave men. One lower jaw would bear an impact mark of a hammer stone. We did not find this trace on the lower jaws of the horizon. Horizon 4 is with a total NISP of 135 and eleven different skeletal elements (apart from the teeth) this horizon the second most diverse one regarding cave hyena material. Hyena coprolites were also collected. According to Dupont (1906) some of the hyena bones were found “vers l’entrée de la caverne a partir de B jusque vers la longer de 60 mètres”; we have not found out what is precisely meant by this statement. Anyhow, Dupont identified nine adults. Also in chamber C numerous hyena bones were found including a lot of juvenile ones. According to Dupont these belonged to seven individuals. This makes a total of 16 hyena individuals found in horizon 4. We only have a total MNI of 13, so maybe some elements are missing.

We see that the third and fourth bone horizon have the most skeletal elements. This horizon lied much deeper than the first two horizons (see 3.1); the bones in this horizon may have been better ‘protected’ from influences like weathering and disturbance. The hyena bones from horizon 4, were found in chamber B and C, which lie the furthest from the entrance. This could again have enhanced preservation.

WALSIN

In this cave 16 different species were found, including the cave hyena. According to Dupont (1906) the cave hyenas occupied the cave as a den; the other animal bones are prey remains. The cave has a total NISP of 73 and eight different skeletal elements, apart from teeth, were found (fig. 4.2B). Duponts work on “L'homme pendant les âges de la pierre dans les environs de Dinant-sur-Meuse” (1875) and his unpublished notes (1906) are the only available sources of information about the excavation of Walsin. Therefore information on this cave is rather limited.

HASTIÈRE

Horizon 1 has a total NISP of 52 (table 4.8) and is characterized by six different skeletal elements apart from the teeth (fig. 4.5A). Dupont (1906) identified five hyena individuals; this corresponds with our calculated MNI (table 4.6). He further mentions that these hyenas were killed by cave men as there are traces of ‘un percuteur’. Horizon 2 harboured bones of nine hyena individuals (our calculated MNI was 13) and some coprolites. Also food remains of cave men were found. This horizon, with ten different skeletal elements, yielded the most diverse hyena assemblage of this cave (apart from the teeth) (fig. 4.5B). Horizon 3 is mainly characterised by the presence of prehistoric men: the proof of their presence consists of modified bones and artefacts.

Specific information about the fourth horizon is lacking as we had no notes of Dupont. Information on horizon 5 is also not available. We found a total NISP of nine and counted a MNI of two. Only four different skeletal elements were counted apart from the teeth and all have about the same frequency (fig. 4.5C). The only research so far on the Hastière material was the study by Dupont in his unpublished notes (1906) and the research of Ehrenberg (1935).

In general, all hyena assemblages found in the three caves all belong to the Age of the Mammoth. The Goyet cave and Hastière cave were not used only by hyenas, this in contrast to the Walsin cave. This latter was according to Dupont (1906) used only by the cave hyena. We cannot deny this assumption, as only the hyena bones were studied but maybe other carnivores used this cave also. The fact that prehistoric men did not occupy this cave is far more certain. As with the words of Dupont (1875) about Walsin: “On constate, au premier coup d’oeil, le contraste entre les ossements recueillis dans une telle caverne et ceux provenant d’une habituation de l’homme”. Dupont refers to the fact that many bones lack their epiphyses but still have their diaphysis and that they show marks of carnivore teeth. Epiphyses are spongy, quite soft and are eaten by a carnivore such as the hyena, leaving the hard diaphysis. Humans, on the contrary, break up the diaphysis with the aid of tools to extract the bone marrow.

5.2 Age determination – Use as communal dens?

In the introduction we refer to the fact that many European caves were visited by cave hyenas and that many of these caves were used as dens. As hyena cubs have an extended juvenile dependency of up to two years (KRUUK, 1972; HOLEKAMP & SMALE, 1998), the presence of cubs among hyena remains are a reliable indication of den occupations in caves (STINER, 2004). Also high numbers and concentrations of coprolites can indicate long-term den occupation. Hyena cubs are born with fully erupted deciduous canine and incisor teeth (KRUUK, 1972; HOLEKAMP & SMALE, 1998). The deciduous cheek teeth start to erupt at two months and the adult teeth are completely erupted at approximately 12-15 months; beginning with the permanent incisors, than the carnassials and finally the premolars and canines (KRUUK, 1972; BINDER & VALKENBURGH, 2000 , VAN HORN et al., 2003). The wear stage III of Stiner (2004) (fig. 3.1) corresponds with an age of approximately 12-15 months. At that time the juvenile hyenas start to leave the communal den (HOLEKAMP & SMALE, 1998).

Deciduous incisors or canine teeth were only found in Goyet (photo 4.4); deciduous cheek teeth were collected in Goyet and Hastière (tables 4.13-4.14, fig. 4.11). Besides these deciduous teeth we also found juvenile jaws (with comparable dentition) in Goyet and Hastière but not in Walsin (fig. 4.12). On the other hand, juvenile permanent incisors and premolars were found in all caves.

Looking at the other age classes, we see that Walsin has the highest number of prime adults (fig. 4.11-4.12). In Goyet and Hastière we see more even age distributions as almost every wear stage is represented. Goyet has with the presence of carnassials germs, the youngest individuals (fig. 4.11 A).

The forgoing data suggest that the cave of Goyet and Hastière were used as communal dens for several reasons. The first factor is the presence of deciduous cheek teeth: they erupt at two months and at this time the cubs reside in the communal den. A second factor is the presence of juvenile jaws, incisors, premolars and molars and juvenile permanent that teeth have a wear- stage of III (= just erupted). Then there is also the presence of germs in Goyet, and in Hastière we have molars with a wear stage II (= erupting). The replacement of the deciduous teeth by the permanent dentition starts at about 12 months and lasts to the age of 15 months. Hyena cubs reside in the communal den till they are about 14 months old. All these factors lead to the conclusion that there were cubs whose age corresponds with the age cubs have when they live in a communal den. Others had the age at which cubs just start to leave a communal den. Then we also found coprolites in both caves, especially Hastière had a relatively high number (24 coprolites). So both ‘conditions’, that is, presence of juvenile remains and coprolites, proposed by Stiner (2004) are present and we can be relatively certain that the caves of Goyet and Hastière were used as communal dens. In Walsin we found no deciduous teeth, only erupting or already erupted permanent teeth. There are also few postcranial juvenile remains present (table 4.2). Coprolites were also not found in this cave so we presume that the Walsin cave was not frequently inhabited or used as den. The age distribution of the M1 (fig. 4.11 B) and the graphs of the jaw per age classes (Fig. 4.12 B) seems to support this conclusion. In the previous section we already mentioned that it is not certain that the cave was inhabited only by cave hyenas. Research on the other bones found in this cave could possibly give clear answers.

We also would like to remark that the excavation methods at the time of Dupont were not so ‘sophisticated’ as today. Small material, like deciduous teeth, were often neglected as the material was seldom or not sieved. Nowadays one uses sieves to retain all small pieces. If the Goyet, Walsin and Hastière caves had been excavated today, we probably would have more small material like deciduous teeth and this would have probably had its effect on our conclusions.

5.3 Gnawing traces and cannibalism

In the introduction we mentioned that cannibalism among hyenas was a somewhat controversial issue with Kurtén (1965) and Kruuk (1972) giving opposite meanings. Today cannibalism among hyenas is widely accepted among both recent and Pleistocene relatives (DIEDRICH & ŽÁK, 2006). Kruuk (1972) mentions that adult hyenas killed in clan fights, or by lions or by other means are eaten by their fellow clan members (although they prefer the meat of their regular prey species). Infants are vulnerable to predation and infanticidal attacks from adult clan members of both sexes (KRUUK, 1972; HOLEKAMP & SMALE, 1998). Cannibalism is not only restricted to adult hyenas: siblicide among hyena occurs frequent. According to Frank (1994) spotted hyenas produce infants that appear to be genetically programmed to attack and kill their siblings. Access to food may be an explaining factor: when food is scarce the dominant cubs kills their subordinates (Frank, 1994; Hofer and East, 2008).

Yet we have to keep in mind that juveniles spend a great deal of time chewing on bones they find in and around the den (HOLEKAMP & SMALE, 1998). It is also not unlikely that adult hyenas will gnaw on bones lying around the den. We therefore would like to make a distinction between intentional and unintentional cannibalism among the hyenas: Intentional cannibalism refers to the killing and eating of living hyenas; with unintentional cannibalism we refer to gnawing on already dead hyenas.

Stiner (2004) states that evidence for interspecific aggression and cannibalism is not easily distinguished in den faunas but that gnawing damage on young juvenile bones is likely to have been from conspecifics, including sibling cannibalism. Diedrick and Žák (2006) state that cracked and chewed bones indicate cannibalism. Diedrich (2006) states that cannibalistic hyenas typically break off the lower jaws and damage the jugals. In general, many authors link hyena gnawing traces with cannibalism, yet they make no distinction between intentional and unintentional cannibalism.

In the assemblages of Goyet, Walsin and Hastière we found hyena bones with gnawing traces of hyenas and most of them belonged to juveniles and sub-adults. (fig. 4.14). The higher frequency of juvenile and sub-adult bones could point to siblicide which makes it likely that intentional cannibalism occurred. Yet again, we cannot be 100% certain as these juveniles may have also died for other reasons and then unintentional cannibalism may have occurred.

5.4 Fossil and recent spotted hyena

The results from our statistical analysis show that there are no significant differences between the fossil hyenas found in Goyet, Walsin and Hastière. In table 5.2 we have summarized some dental measurements of Goyet, Walsin and Hastière and of the Spy cave (GERMONPRÉ et al., in press) and the Marie-Jeanne cave (GAUTIER & HEINZELIN, 1980). All these measurements are compared with the results found by Kurten on British fossil hyenas (1965). Except for Walsin, the lengths of the lower carnassials (M1) are more than 32 mm, indicating the hyena material date from the Last Glacial (KLEIN & SCOTT, 1989; GERMONPRÉ et al., in press). The measurements from Goyet, Walsin and Hastière fall within the values of Spy, Marie-Jeanne and Great-Britain which could indicate that these hyenas belonged to the same European population.

Mean Goyet Walsin Hastière Spy Marie-Jeanne Weichselian* Eemian* P3 cl 24.48 23.55 23.34 24.01 24.39 24.72 23.70 P3 cw 17.96 17.42 17.11 17.68 17.40 18.33 18.32 P4 cl 40.02 42.56 41.70 41.61 40.76 - - P4 cw 21.56 20.83 21.41 22.32 - - - P2 cl 16.68 16.42 16.50 16.25 16.93 16.42-16.72 17.22-17.50 P3 cl 22.63 22.83 22.40 22.39 22.60 22.43-22.94 21.52-21.75 M1 cl 32.24 31.40 32.80 32.41 32.37 - - M1 cw 14.00 14.11 13.97 14.10 13.77 13.86-14.30 13.38-13.57

Table 5.2: Average dental measurements of Goyet, Walsin, Hastière, Spy (GERMONPRÉ et al., in press) and Marie-Jeanne (GAUTIER & HEINZELIN, 1980) - * Kurten (1965)

Between the fossil spotted hyena and the recent spotted hyena we found significant differences (table 4.62-4.63). The scatter plot of the lower P2 and P3 (fig. 4.20 – 4.21) show that these fossil hyena teeth are larger than the recent ones. The scatter plot in fig. 4.22 shows less difference between the two skulls of the fossil hyenas and those of the recent hyenas.

Differences between fossil cave hyenas and recent spotted hyenas were described by Kurten (1965); and they are especially morphological Fossil cave hyenas are bigger, have a slightly different cranial morphology and show differences in jaws and teeth (KURTÉN, 1965; ROHLAND et al., 2005). Klein and Scott (1989) found glacial/interglacial size variations among British fossil spotted hyenas; they concluded that spotted hyena body size is inversely related to temperature as predicted by Bergmann’s rule. Meiri et al. (2004) showed that the occurrence of Bergmann’s rule in the Carnivora is less frequent than earlier published data suggest but the spotted hyena would follow indeed the rule of Bergmann.

6. CONCLUSION

The fossil hyena assemblages found in Goyet, Walsin and Hastière originated from the same European cave hyena population as the cave hyenas from Spy, Marie-Jeanne and Great-Britain. This population lived during the Last Glacial.

The caves of Goyet and Hastière were not only visited and inhabited by cave hyenas, other carnivores and prehistoric man occupied these caves also. Furthermore, both caves were used as communal dens; this is reflected by their hyena assemblages (presence of juvenile remains).

The Walsin cave was visited by cave hyena, but it is not clear from the hyena assemblage if they used it as communal den. It is also possible that other carnivores used the cave but future research is needed to obtain clear answers.

Whether cannibalism among the cave hyenas from Goyet, Walsin and Hastière occurred, is not straightforward. As mentioned in section 5.3, a difference between intentional and unintentional cannibalism should be made. In our study, we cannot conclude which form of cannibalism occurred. The high frequency of juvenile bones may indicate that siblicide, a behavioural pattern that occurs frequently among the recent spotted hyenas, also occurred among the cave hyenas.

The cave hyenas or fossil spotted hyenas are bigger than the recent spotted hyenas and this is consistent with the literature (KURTÉN, 1965, KLEIN & SCOTT, 1989, ROHLAND ET AL., 2005).

7. DUTCH SUMMARY

In deze masterproef werden de fossiele resten van de holenhyenas uit drie Belgische grotten, namelijk de grotten van Goyet, Walsin en Hastière uit de provincie Namen, geanalyseerd. Al het fossiele materiaal behoort tot de Dupont collectie, genoemd naar E. Dupont die deze grotten uitgroef tussen 1866 en1872. Het materiaal wordt bewaard op het Departement Paleontologie van het Koninklijk Belgisch Instituut voor Natuurwetenschappen (K.B.I.N.) Eén fossiele schedel, die werd gebruikt in dit onderzoek, komt uit de Noordzee en behoort tot de fossiele collectie van de Paleontologische Unit van de Universiteit Gent.

Rohland et al. (2005) toonden aan dat de fossiele holenhyena en de recente Afrikaanse gevlekte hyena nauw verwant zijn aan elkaar. We kunnen dus gebruik maken van de kennis over de levenswijze van de recente gevlekte hyena om interpretaties te maken over het leven van deze fossiele holenhyenas. De jongen van recente gevlekte hyena’s worden geboren in een afgelegen werphol. Hun ogen zijn dan al volledig open en ze bezitten reeds snij-en hoektanden. Na ongeveer een maand worden de jongen overgebracht naar een gemeenschappelijk hol waar ze verblijven tot ze ongeveer negen maanden oud zijn.

In dit onderzoek trachtten we na te gaan of de grotten van Goyet, Walsin en Hastière gebruikt werden door de holenhyenas als gemeenschappelijk hol voor hun jongen. De aanwezigheid van juveniel hyena materiaal is daarvan een goede aanwijzing. Daarnaast trachtten we ook na te gaan of kannibalisme voorkwam. Tot slot vergeleken we het fossiel materiaal met wat recent materiaal. Dit recente hyena materiaal wordt bewaard in het KBIN op het Vertebraten Departement.

Bijna al het hyenamateriaal werd geanalyseerd en opgemeten. Een belangrijke stap hierbij was het bepalen van leeftijdsklassen. Voor de individuele en kaaktanden maakten we gebruik van de tabel van Stiner (2004) over het doorbreken en de slijtage van tanden. Op die manier werd het craniaal materiaal voorzien van drie leeftijdsklassen (“juvenile”, “prime adult”, “old adult”). Voor het postcraniale materiaal keken we naar de vergroeiing van de epifyse; dit leidde ook tot een verdeling in drie leeftijdsklassen (juvenile, sub-adult, adult). De leeftijdsklassen van het craniaal en postcraniaal materiaal komen niet geheel overeen, maar het postcraniale ‘leeftijdsdateren’ werd hoofdzakelijk gedaan om de juvenielen te onderscheiden. Ook voerden we enkele statistische testen uit om na te gaan of er significante verschillen waren tussen de fossiele holenhyenas en de recente gevlekte hyena’s.

Uit het onderzoek blijkt dat de grotten van Goyet en Hastière gebruikt werden als gemeenschappelijk hol. Zo vonden we enkele melktanden en een aantal juveniele onderkaken. De leeftijdsverdelingen van deze twee grotten komen overeen met het tijdstip waarop de jongen nog verbleven in het gemeenschappelijk hol en het hol net begonnen te verlaten. Ook vonden

73 we in deze twee grotten een aantal coprolieten terug. Het terugvinden van juveniel materiaal en coprolieten stemt overeen met de bevindingen van Stiner (2004) aangaande de bezetting van een hol door hyena’s. In de grot van Walsin vonden we noch melktanden noch juveniele kaken terug. Er was wel wat juveniel postcraniaal materiaal aanwezig. Uit de leeftijdsverdeling blijkt dat de grot voornamelijk werd bezet door ‘prime adults’ en ‘old adults’. Voor deze grot kunnen we niet met 100% zekerheid besluiten dat ze werd gebruikt als gemeenschappelijk hol.

Uit de tafonomie van de grotten bleek verder dat zowel de grot van Goyet als de grot van Hastière ook gebruikt werden door andere carnivoren (zoals de holenbeer) en door prehistorische mensen. De grot van Walsin werd niet bezet door prehistorisch mensen; een gebruik door andere carnivoren is niet uit te sluiten. Daarvoor is verder onderzoek nodig op het andere faunale materiaal van deze grot.

Op enkele hyenabeenderen werden knaagsporen gevonden van hyena’s. In veel literatuur worden knaagsporen van hyena’s op hyenabotten gezien als kannibalisme. Onderzoek naar de levenswijze van de recente gevlekte hyena leert ons dat hyena’s elkaar opzettelijk kunnen aanvallen en opeten. Het gaat dan vooral om adulte mannetjes die jongen aanvallen of over siblicide. Onderzoek en studie van de gevlekte hyena heeft echter ook aangetoond dat de jongen knagen op alle beenderen die zij kunnen vinden in (of in de omgeving van) het gemeenschappelijk hol. Ook adulte hyena’s knagen nu en dan eens op rondslingerende beenderen. Knaagsporen zijn dus niet alleen afkomstig van opzettelijke kannibalistische acties. Daarom ijveren we voor een onderscheid tussen het opzettelijk doden van andere hyena’s (“intentional cannibalism”) en het beknagen van door andere oorzaken omgekomen hyena’s (“unintentional cannibalism”). Er was dus kannibalisme maar we kunnen verder niet specificeren over welke vorm van kannibalisme het in deze gevallen gaat.

We vonden geen significante verschillen tussen de tandmetingen van de fossiele hyena’s van Goyet, Walsin en Hastière. Daarnaast vergeleken we het fossiele tandmateriaal van deze drie grotten met de tandmetingen van Spy, Marie-Jeanne en Groot-Brittannië. Hieruit blijkt dat er geen regionale verschillen waren en dat de hyena’s behoorden tot dezelfde Europese populatie.

De vergelijking van het fossiele en recente tand-materiaal gaven wel significante verschillen. Uit een scatterplot met kroonlengtes en kroonbreedtes van de P2’s en P3’s blijkt dat de deze tanden bij de fossiele hyena’s groter waren dan bij hun recente verwanten. Deze bevindingen sluiten aan bij de literatuur (KURTÉN, 1965, KLEIN & SCOTT, 1989, ROHLAND et al., 2005).

8. THANKS

I would like to thank some people who contributed to this thesis:

Firstly, I want to thank Dr. M. Germonpré for the excellent accompaniment. She was always prepared to answer my questions, she gave my tips and aided in the search for literature. I am also grateful for the time she has spent in correcting my texts.

I want to thank Prof. Dr. A. Gautier for his corrections and advices concerning my texts. I also want to thank him for readying a fossil hyena skull.

I am Prof. Dr. J. Verniers thankful for his promotorship; he contacted Dr. M. Germonpré which made this thesis possible.

The Belgian Royal Institute for Natural Sciences owes my gratitude for the provision of the material and for taking the photographs.

Finally, I would like to thank my family, friends and all other persons who supported me while writing this thesis, but I also would like to thank them for their support during my five-year biology study.

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ANNEX

Alphabetical list of abbreviations ad.: adult C: Canine carp: carpal(s) d: decidious dm: deciduous (pre)molar di: deciduous incisor fig.: figure I: Incisor ID: Identification number ind. or indet.: indeterminable juv.: juvenile M: Molar max.: maxilla (=upper jaw) mand.: mandibula (=lower jaw) MC: metacarpal(s) MT: metatarsal(s) n: total number number DP: number from Dupont P: Premolar Photo: Photograph pr. ad.: prime adult o.ad.: old adult Stv dev: Standard deviation subad.: subadult tars: tarsal(s)

Statistical tests: on added CD-Rom

Primary data: on added CD-Rom