Crocuta spelaea Goldfuss, 1823

Sauqué, V, Rabal-Garcés, R, Madurell-Malaperia, J, Gisbert, M, Zamora, S, Torres, T, Ortíz, J & Cuenca-Bescós, G, 2017, Pleistocene cave hyenas in the Iberian Peninsula: New insights from Los Aprendices cave (Moncayo, Zaragoza, Palaeontologia Electronica (Ankara, Turkey) 20 (1), pp. 1-38 : 7-19

Crocuta spelaea Goldfuss, 1823

( Figures 4-7 )

1823 Hyaena spelaea Goldfuss p.456-462, figs. 16, 17.

1993 Crocuta crocuta spelaea Cardoso , p. 82-99, figs. 1, 2, 3, 4.

v. 1997 Crocuta crocuta Blasco and Montes , p.12, 14, 17-21, pl, 2, 3, 4, 5, 6, 7.

1998 Crocuta crocuta spelaea Iñigo, Molero and Maldonado , p. 66, 68, fig. 1.

1999 Crocuta spelaea Baryshnikov , p. 167, 168, figs. 2, 3, 4.

2002 Crocuta crocuta Davis , p. 54, 55, fig. 21.

2005 Crocuta crocuta Arbizu, Álvarez-Lao, and Adan , p. 134, pl, 1..

2006 Crocuta crocuta spelaea Diedrich and Žák , p. 248, 249, figs. 5, 6.

2006 Crocuta crocuta spelaea Testu , p. 273, 274, pl, 12, 13

2008 Crocuta crocuta spelaea Diedrich , p. 825, fig. 3.

2010 Crocuta crocuta Arsuaga, Baquedano, Pérez-González, Sala, García, Álvarez-Lao, Laplana, Huguet, Sevilla, Blain, Quam, Ruiz Zapata, Sala, Gil García, Uzquiano, and Pantoja , p. 63, fig. 9.

2010 Crocuta spelaea Rosell, Blasco, Rivals, Cebrià, Morales, Rodríguez, Serrat, and Carbonell , fig. 4.

2012a Crocuta crocuta spelaea Diedrich , p. 67-74, figs. 3, 4, 5, 6, 7, 8, 9,10.

2012b Crocuta crocuta spelaea Diedrich , p. 173-177, figs. 3, 4, 5.,6,7

v. 2012 Crocuta crocuta Martínez-Sánchez, López-García, Alcalá-Ortiz, Blain, and Rabal-Garcés , p. 551, fig. 2.

2014 Crocuta crocuta Maroto , fig. 4.

2015 Crocuta crocuta Álvarez-Lao, Ruiz-Zapata, Gil-García, Ballesteros, and Jiménez-Sánchez , fig. 5.

2015 Crocuta crocuta spelaea Fourvel, Fosse, Fernandez, and Antoine , p. 248, 249, 250, fig. 3.

TABLE 1. List of anatomical elements recovered from the individual of Crocuta spelaea from Los Aprendices. L left. R right. NR, number of remains; NISP, number of identified remains; MNE: minimum number of elements.

Elements MNE NISP NR

Cranium 1 12 12

Mandible R 1 1 1

Mandible L 1 3 3

Axis 1 4 4

CervicalV. 5 9 9

Thoracic V. 7 26 26

Lumbar V. 4 8 8

Ribs 10 29 29

Scapula L 1 2 2

Scapula R 1 2 2

Humerus R 1 2 2

Ulna L 1 2 2

Ulna R 1 4 4

Radius R 1 3 3

Scapholunate R 1 1 1

Capitate R 1 1 1

Unciform R 1 1 1

Capitate L 1 1 1

Pisiform R 1 1 1

Mtc II R 1 1 1

Mtc III R 1 1 1

Mtc IV R 1 1 1

MtcV R 1 1 1

Mtc II L 1 1 1

Mtc III L 1 1 1

Holotype. Hyaena spelaea Goldfuss, 1823 . Incomplete cranium (Goldfuss-Museum Bonn No. M2609).

Type Locality. Zoolithen Cave, Geilenreuth, Wiensent Valley, Bavaria, Germany.

Localities. The species is found in hundreds of Pleistocene caves throughout Europe.

Material. cranium ( MPZ 2014/657), one mandible ( MPZ 2014/593), one axis ( MPZ 2014/589), five cervical vertebrae ( MPZ 2014/592, MPZ 2014/605, MPZ 2014/607, MPZ 2014/668, MPZ 2014/675), seven thoracic vertebrae ( MPZ 2014/641, MPZ 2014/656, MPZ 2014/661, MPZ 2014/667, MPZ 2014/670, MPZ 2014/674, MPZ 2014/690), four lumbar vertebrae ( MPZ 2014/582, MPZ 2014/586, MPZ 2014/701, MPZ 2014/703), ten ribs ( MPZ 2014/591, MPZ 2014/604, MPZ 2014/612, MPZ 2014/637, MPZ 2014/646, MPZ 2014/651, MPZ 2014/652, MPZ 2014/658, MPZ 2014/662, MPZ 2014/666), two scapulae ( MPZ 2014/584, MPZ 2014/588), one humerus ( MPZ 2014/672), two ulnae ( MPZ 2014/610; MPZ 2014/676), one radius ( MPZ 2014/611), one scapholunate ( MPZ 2014/ 613), two capitate ( MPZ 2014/630, MPZ 2014/659) one pisiform ( MPZ 2014/614), eight metacarpi ( MPZ 2014/575, MPZ 2014/581, MPZ 2014/615, MPZ 2014/616, MPZ 2014/617, MPZ 2014/618, MPZ 2014/684, MPZ 2014/685), one femur ( MPZ 2014/587), two tibiae ( MPZ 2014/671, MPZ 2014/ 681), one talus ( MPZ 2014/682), two calcanei ( MPZ 2014/632, MPZ 2014/677), one unciform ( MPZ 2014/678), two cuboids ( MPZ 2014/ 623, MPZ 2014/ 629), two scaphoids ( MPZ 2014/623, MPZ 2014/629), two cuneiforms ( MPZ 2014/62, MPZ 2014/628), eight metatarsi ( MPZ 2014/633, MPZ 2014/634, MPZ 2014/635, MPZ 2014/636, MPZ 2014/642, MPZ 2014/643, MPZ 2014/644, MPZ 2014/645) and twenty-four phalanxes ( MPZ 2014/575, MPZ 2014/576, MPZ 2014/577, MPZ 2014/578, MPZ 2014/579, MPZ 2014/580, MPZ 2014/583, MPZ 2014/590, MPZ 2014/594, MPZ 2014/599, MPZ 2014/600, MPZ 2014/601, MPZ 2014/602, MPZ 2014/603, MPZ 2014/620, MPZ 2014/621, MPZ 2014/622, MPZ 2014/626, MPZ 2014/627, MPZ 2014/631, MPZ 2014/639, MPZ 2014/654, MPZ 2014/655, MPZ 2014/680).

The cave hyena is the predominant taxon in Los Aprendices. As the elements were found in

SAUQUÉ ET AL.: CROCUTA SPELAEA IN IBERIA anatomical position and, presumably because there is no duplication of elements, they have been assigned to a single individual. We here describe the cranium, mandible, teeth, because these are elements of great taxonomic importance.

Description

Cranium. The cranium (MPZ 2014/657) is almost complete, lacking a fragment of the right zygomatic arch and part of the nasals including a fragment of the right side. The sagittal crest is slightly deformed, and the left side of the neurocranium is flattened ( Figure 5 ). The diastema between the canine and P1 is reduced, and P1 and P2 are in contact ( Figure 5.4 ). In extant Crocuta crocuta , P1- P2 are separated by a distance of several mm. The infraorbital foramen is located above the point of contact between P2 and P3. The dental series displays the moderate curvature typical of C. spelaea ( García, 2003) . The palate is wide at the P4, and even though it is broken a pronounced concavity characteristic of the genus Crocuta can be observed (Morales et al., 1987). The cranium has a wide snout with a slight post-canine constriction. The rostral region (splanchnocranium) is short and wide, conferring upon it a “U”-shape in dorsal view, whereas in Hyaena it is “V”-shaped ( García, 2003). One of the most notable characteristics of the cranium is the absence of M1. This element is generally lacking in the genus Crocuta but is present in Hyaena, in which M1 has three cuspids ( Kurtén, 1957; Werdelin and Solounias, 1991; García, 2003).

The cranium has a total length of 305 mm, which lies clearly outside the range of extant Crocuta crocuta (see Figures 5 , 8 ; Appendix 2). However, it matches the size documented for cave hyenas and is larger than the C. c. intermedia ( Serres et al., 1828) of Lunel Viel ( Bonifay, 1971). In general terms the measurements of the skull of Los Aprendices are similar but lightly larger than those of Crocuta crocuta praespelaea from Mosbach, Kazanka and Petralona ( Kurtén, 1962; Kurtén and Poulianos, 1981; Schütt, 1971; Baryshnikov, 2014). The specimen from Los Aprendices is similar in size to cave hyenas from central Europe at the sites of Badel Cave, the Perick Caves, and Rösenbecker Cave ( Diedrich, 2011a), as well as at the French site of Jaurens ( Ballesio, 1979). In comparison with other crania from the Iberian Peninsula, the specimen from Los Aprendices is clearly smaller as compared with the specimen from Fontainhas (L= 317 mm) in Portugal ( Cardoso, 1993), and slightly larger than the cranium from Gabasa 1 (L= 294mm) ( Blasco and Montes, 1997). Furthermore, it is larger than the latest Early Pleistocene cranium from Gran Dolina TD4W (L= 265mm), which is ascribed to C. crocuta ( García and Arsuaga, 1999) .

Upper dentition. C1. The length and width of the upper canines from Los Aprendices are within the range of variation of extant Crocuta crocuta but are clearly larger than average (Appendix 3).

P1. P1 is a small simple tooth, the crown of which forms a low cone crossed by a ridge that transversely divides it into two.

P2. Mesial and distal accessory cusps are present and clearly developed. The outline of the tooth is rectangular, with the distal part buccolingually wider than the mesial part. The length and width of P2 from Los Aprendices fall within the range of extant Crocuta crocuta but close to the maximum values. The P2 from Los Aprendices is clearly larger than the P2 from the cranium of C. crocuta from Gran Dolina TD 4W. The P2 from Los Aprendices is larger than C. crocuta spp. from Petralona ( Baryshnikov and Tsoukala, 2010) and the C. c praespelaea from Kazanka and Mosbach ( Baryshnikov, 2014), which according to Baryshnikov (2014) has the largest P2 of the spelaea forms. It is remarkable that the Los Aprendices P2 is the largest premolar recovered in the Iberian Peninsula of fossil record of Crocuta genus.

P3. P3 is characterized by the great development of the protocone, which is surrounded by a well-developed cingulum and lacks accessory cusps. This tooth is an important part of the process of bone crushing (Schütt, 1971), and therefore its structure can be used to evaluate the degree of adaptation the dentition to feeding on carrion ( Baryshnikov, 2014). This tooth presents an extremely high crown, which is also seen in other populations of Crocuta spelaea . It indicates a Crocuta spelaea specialization for breaking bones related to scavenging. The length and width values of P3 from Los Aprendices are within the range of variation of the Iberian Pleistocene cave hyena and exceed the measurements of extant C. crocuta (Appendix 3).

P4. The anterior face of the protocone is approximately level with the anterior face of the parastyle, which is typical of the genus Crocuta (Wederlin and Solounias, 1991) Its greatest length and width markedly exceed those of extant C. crocuta and C. c. intermedia ( Cardoso, 1993), and it is also larger than the C. crocuta from the Middle Pleistocene of Gran Dolina TD4W ( García, 2003). The Los Aprendices specimen is of similar length and width as C. c. praespelaea from Kazanka ( Baryshnikov, 2014) and is slighty larger than C. c. praespelaea from Mosbach (Bayshnikov, 2014) and C. crocuta spp. from Petralona ( Baryshnikov and Tsoukala, 2010). Therefore, its values are similar to those of C. spelaea from the Iberian Peninsula, such as Gabasa 1 ( Blasco and Montes, 1997), Fontainhas ( Cardoso, 1993), Sima I, Kobaedaerra ( Castaños, 1987), Labeko Koba ( Altuna and Mariezkurrena, 2000), El Toll, Olopte B and Cova de l’Or (J.M.-M personal commun., 2015.) (Appendix 3). The extreme metrical variability of dental remains ( Figure 9 , Appendix 3) most likely coincides with ecomorphotypic variation, so it is not recommended to use cave hyenas for biochronological purposes (e.g., Kurtén, 1957; Kurtén and Poulianos 1977; Klein and Scott 1989; Baryshnikov 1999, Fourvel et al., 2015).

Mandible. The left hemimandible from Los Aprendices is well preserved and practically complete. The corpus preserves the i1, i2, c1, p2, p3, p4 and m1, all showing wear. The ramus has a complete and well preserved coronoid process, articular and angular apophyses. The corpus presents a basal edge that in lateral view is strongly convex at the level of m1 ( Figure 5.5-6 ). This feature is characteristic of Crocuta ( Bonifay, 1971; Testu, 2006). The hemimandible is mesiolingually curved. In buccal view, the ramus in its distal part shows a deep, subtriangular masseteric fossa that extends from the protoconid of m1 to the angular process and the condyle. The single mental foramen is placed at the level of p2. The arrangement of the first two premolars (p2 and p3) is not overlapping, and p4 and m1 overlap as in Crocuta (Testu, 2006) . The disposition of the premolars in the dental series is clearly curved ( Figure 5.7 ); this curvature is caused by the increase in size and is a distinctive feature of C. spelaea ( García, 2003) .

Los Aprendices hemimandible is larger mesiodistally as compared with extant Crocuta crocuta (Appendix 4). It is also far larger than fossil hyenas from the Middle Pleistocene, such as Atapuerca TD 8 ( García, 2003) and Lunel Viel ( Bonifay, 1971). However, the distance c1-m1 is smaller than in C. c. praespelaea from Kazanka ( Baryshnikov, 2014). On the other hand, the mandible fits perfectly within the range of C. spelaea and is one of the largest specimens of the Iberian Peninsula (Appendix 4). The biometric analysis of the hemimandible further reinforces the inclusion of the studied specimen in the hypodigm of C. spelaea .

Lower dentition. c1. The c1 is within the range of variation shown by the population of Pleistocene cave hyenas and larger than extant Crocuta crocuta (Appendix 5).

p2. The main cusp, the protoconid, is buccolingually wide and mesially surrounded by a well-developed cingulum. The distal accessory cusp is well-developed and separated from the protoconid by a transverse groove. The p2 of Los Aprendices is clearly larger than the average for extant C. crocuta , C. c. intermedia and C. c. praespelaea from Petralona (Appendix 5). On the other hand, its values fit perfectly with the C. spelaea from Labeko Koba ( Altuna and Mariezkurrena, 2000) and are slightly smaller than the individual of C. c. praespelaea from Kazanka ( Baryshnikov, 2014). It is also slightly larger than the population of C. spelaea form Gabasa ( Blasco and Montes, 1997).

p3. The protoconid has a slightly distal tilt, and shows a marked cingulum on the mesial and distal edges. No accessory cusps are developed. The p3 of Los Aprendices is the largest from the Iberian fossil record and definitely larger than extant Crocuta crocuta (Appendix 5).

p4. p4 displays well-developed accessory cuspid with the distal one being mesiodistally longer than the mesial one. As in p3, the protoconid displays a slight distal tilt. The p4 fits perfectly within the range of Late Pleistocene cave hyenas, being larger than extant Crocuta crocuta and C. c. intermedia, which we classified as a C. crocuta from the Middle Pleistocene. Furthermore, the measurements from Los Aprendices are similar to the C. c. praespelaea (Appendix 5).

m1. The paraconid is clearly mesiodistally larger than the protoconid, a taxonomic character only seen in Crocuta among Middle to Late Pleistocene bone-cracking hyaenas (Werdelin and Solounias, 1991). The m1 presents a well-developed cingulum on the lingual side that extends up to the anterior edge. Distally, a mesiodistally short talonid is evident, which is a character typical of the genus Crocuta . Finally, the distolingual part of the protoconid displays a small but well-marked metaconid. Several scholars consider the presence of well-developed metaconid in the m1 as an ancestral character in the genus Crocuta , barely present in the last European spotted hyaenas. However, the extreme variability in the development and presence of m1 metaconid in extant and extinct representatives of the genus Crocuta prevent its use for taxonomic proposes (Werdelin and Solounias, 1991; García, 2003).

The length and width of m1 from Los Aprendices are within the range of variation of the Iberian Pleistocene cave hyena and exceed the measurements of extant Crocuta crocuta (Appendix 5). In Figure 10, it can be observed that there are two distinct populations: one for the C. spelaea and another population made up of extant C. crocuta and the specimens of C. crocuta from the Middle Pleistocene, such as C. c. intermedia ( Bonifay, 1971) C. crocuta from Gran Dolina ( García, 2003).

Forelimbs. The forelimb bones ( Figure 6 ; Appendix 6) include both incomplete scapulae, one nearly complete humerus, both complete ulnas, one complete radius, some carpals such as the scapholunar, pisiform, cuneiform and unciform, and all eight metacarpals except the Mtc I.

Humerus. The proximal epiphysis of the humerus is broken and the head is missing. In cranial view, the diaphysis is straight laterally flattened, with caudal bending in the proximal third. The deltoid crest is very prominent and connected with the humeral crest. The distal trochlea is wide and the condyle is broken. Above this there is a well-developed oval supratrochlear foramen. In the distal part of the diaphysis, there is a deep and well-developed olecranon fossa. As a consequence of the broken or absent epiphysis, is not possible to metrically compare the humerus with other known samples.

Radius. The proximal epiphysis has an oval outline. The bicipital tuberosity is prominent, oval and FIGURE 10. Graph representing total length versus total width of m1 of extant and cave hyaena from several sites. The comparative measurements (in mm) have been extracted from, Reynolds, 1902; Bonifay, 1971; Ballésio, 1979; Clot, 1980; Argant, 1991; Cardoso, 1993; Blasco and Montes, 1997; Fosse 1997; Castaños, 1987; Baryshnikov, 1999; Altuna and Mariezkurrena, 2000; García, 2003, Testu, 2006; Fourvel, 2012 and J.M.-M personal commun., 2015. Abbreviations: ATA, Atapuerca; Ap, Los Aprendices; Ar, Arbreda; Fo, Fontainhas; IPS, Institut de Paleontologia M. Crusafont de Sabadell; Mo, Mollet; To, El Toll; Va, Valdegoba. Confidence ellipses with confidence interval (0.95) are figured.

is placed on the proximolateral part of the diaphysis. The diaphysis has a semicircular cross section and is anteroposteriorly flattened. It shows strong torsion in the distal part of the bone. The distal epiphysis is mediolaterally wider than the diaphysis and has a very marked styloid process that is pointed and projected distally. The radius length of the Los Aprendices specimen (227 mm) is within the range of variation of Iberian Pleistocene cave hyenas. Its length also lies within the range of extant Crocuta crocuta , but is slightly below average (Appendix 6, Figure 11.1 ). Despite being shorter than that of extant C. crocuta , it is moderately mediolaterally wider, which gives a characteristic robust aspect to C. spelaea ( Kurtén, 1957) .

Ulna. The proximal part of the olecranon process is not preserved and the diaphysis displays more curvature and robusticity as compared with extant Crocuta crocuta . The triangular area below the lesser semilunar notch is deep and connected to the depressed area for the interosseous ligament, which can be seen on the lateral part of the diaphysis. The styloid process is robust with a rounded articular circumference that projects posteriorly.

Metacarpals. The metacarpals are clearly shorter than the average in extant Crocuta crocuta and are mediolaterally robust (Appendix 6, Figure 11.2 ). The length and width of MtcpIII falls outside the range of variation of extant C. crocuta , being closer to the minimum values for proximodistal length and close to the maximum values for mediolateral breadth (Appendix 6).

Hindlimbs. Of the hindlimbs ( Figure 7 ; Appendix 6), only one broken femur and both tibiae are more or less complete. One astragalus and both calcanei complete some tarsals have also been recovered: one navicular, both cuboids, one first cuneiform and all eight metatarsals except Mtt I.

Femur. The diaphysis is straight in the proximal part and slightly anterioposteriorly curved in the distal part. It has a circular cross-section, which becomes oval in the distal part. The proximal epiphysis is broken and the great trochanter is missing. The neck is complete and connects the diaphysis with the head, which is spherical shape and presents an oval fovea capitis. The distal epiphysis of femur consisted of two similar condyles. The lateral one is partially broken. The condyles are similar, situated at the same level and separated by a intercondyloid fossa. The femur length of the Los Aprendices specimen is within the range of variation of extant Crocuta crocuta but very close to the maximum values and clearly greater than the average (Appendix 6). The distal epiphysis mediolateral width is outside the range of variation of extant C. crocuta (Appendix 6). The femur from Los Aprendices is similar in size to the femur from Labeko Koba ( Altuna and Mariezkurrena, 2000) and larger than the femur from TD5, which belongs to C. crocuta ( García, 2003) . In Figure 12.1 , it can be observed that there are two clusters that differentiate C. spelaea from extant C. crocuta .

Tibia. The tibia is short and robust. It is slightly convex on the proximomedial side. The proximal epiphysis has a triangular cross-section, and this morphology continues throughout the diaphysis except for the distal portion, where it is oval. The tibial crest is wide and well-developed. The distal epiphysis is subrectangular, with a well differentiated, laterally projecting medial rounded malleolus. The length and width of tibia from Los Aprendices are within the range of variation of the cave hyena and extant Crocuta crocuta (Appendix 6, Figure 12.2 ).

Metatarsals. The metatarsals are shorter than those of extant Crocuta crocuta and more robust (Appendix 6, Figure 12.3 ). As occurs with Mtc III, the length and width of Mtt III is outside the range of variation of extant C. crocuta , but close to the length minimum and wider than the maximum value (Appendix 6, Figure 12.3 ).

Figures

FIGURE 4. Panoramic photo of the skeleton of Crocuta spelaea from Los Aprendices.

FIGURE 5. Photograph of the cranium (1-4, MPZ 2014/657) and hemi-mandible (5-7, MPZ 2014/593) of Crocuta spe- laea: 1-2, lateral view; 3, dorsal view; 4, occlusal view; 5, buccal view; 6, lingual view; 7, occlusal view.

FIGURE 6. Crocuta spelaea forelimb remains from Los Aprendices. 1, left scapula in dorsal view (MPZ 2014/584). 2, left humerus in cranial view (MPZ 2014/672). 3, right radius in palmar view (MPZ 2014/611). 4, left ulna in medial view (MPZ 2014/610). 5, right ulna in medial view (MPZ 2014/676). 6, right Mtc V in dorsal view (MPZ 2014/618). 7, right Mtc IV in dorsal view (MPZ 2014/617). 8, right Mtc III in dorsal view (MPZ 2014/616), 9, right Mtc II in dorsal view (MPZ 2014/615). 9, left Mtc II in dorsal view (MPZ 2014/685). 10, left Mtc III in dorsal view (MPZ 2014/684). 11, left Mtc IV in dorsal view (MPZ 2014/ 581). 12, left Mtc V in dorsal view (MPZ 2014/575). 13, right scapholunate in proximal view (MPZ 2014/613). 14, right pisiform in proximal view (MPZ 2014/614). 15, trapezoid (MPZ 2014/619). 16, right cuneiform in dorsal view (MPZ 2014/659). 17, unciform (MPZ 2014/678) 18-26, first phalanx in dorsal view (MPZ 2014/576, 577, 578, 583, 621, 622, 626, 631). 27-34, second phalanx in lateral view (MPZ 2014/594, 599, 600, 601, 627, 639, 654, 655). 35-37, third phalanx in dorsal vier (MPZ 2014/590, 603, 680).

FIGURE 7. Crocuta spelaea hind limb remains from Los Aprendices. 1, right femur in caudal view (MPZ 2014/587). 2, left tibia in cranial view (MPZ 2014/681). 3, right tibia in cranial view (MPZ 2014/671). 4, left calcaneus in dorsal view (MPZ 2014/677). 5, right calcaneus in dorsal view (MPZ 2014/632). 6, left astragalus in dorsal view (MPZ 2014/682). 7, left navicular in lateral view (MPZ 2014/625). 8, right cuboid in proximal view (MPZ 2014/ 623). 9, left cuboid in proximal view (MPZ 2014/ 629). 10, left first cuneiform in proximal view (MPZ 2014/62). 11-14, first phalanx in dorsal view (MPZ 2014/579, 580, 620, 624). 15, right Mtt V in dorsal view (MPZ 2014/636). 16, right Mtt IV in dorsal view (MPZ 2014/635). 17, right Mtt III in dorsal view (MPZ 2014/634). 18, right Mtt II in dorsal view (MPZ 2014/633). 19, left Mtt II in dorsal view (MPZ 2014/642). 20, left Mtt III in dorsal view (MPZ 2014/643). 21, left Mtt IV in dorsal view (MPZ 2014/ 644). 22, left Mtt V in dorsal view (MPZ 2014/645).

FIGURE 8. Bivariate graph representing the width of the condyles in relation to the total length of the cranium. Abbreviations:Ba, Badel Cave; BW, Bad Wildungen; Do, Döbritzer Cave; Ja, Jaurens; Ke, Ketsch; Pe, Perick Caves; Ro, Rösenbecker Cave; Th, Thiede; Oa, Oase Cave; Sr, Srbsko Cave; Vy, Výpustek Cave (Diedrich, 2011a); Sl, Sloup Cave (Diedrich, 2012b); Ga, Gargas (Cardoso, 1993); Lu, Lunel Viel (Bonifay, 1971). The measurements of the crania of extant C. crocuta specimens were taken by J.M.-M of the Royal Museum for Central Africa (RMCA) of Belgium.

FIGURE 9. Graph representing total length versus total width of P4 of extant and cave hyaena from several sites. The comparative measurements have been extracted from, Reynolds, 1902; Bonifay, 1971; Ballésio, 1979; Clot, 1980; Argant, 1991; Cardoso, 1993; Blasco and Montes, 1997; Fosse, 1997; Castaños, 1987; Baryshnikov, 1999; Altuna and Mariezkurrena, 2000; García, 2003; Testu, 2006; Fourvel, 2012; and J.M.-M personal commun., 2015. Abbreviations: ATA, Atapuerca; Ag, Aguilón P-7; Ap, Los Aprendices; Mo, Mollet; Ol, Olopte B; L’Or Cova de l’Or; SI, Sima I. Confidence ellipses with confidence interval (0.95) are figured.

FIGURE 11. 1, Graph representing total length of radius versus distal width of extant and cave hyaena from several sites. The comparative measurements (in mm) have been extracted from Ehrenberg, 1940; Cardoso, 1993; García, 2003; Diedrich, 2011e; and J.M.-M personal commun., 2015. 2, Total length of third metacarpus versus smallest breadth of the diaphysis (SD) of extant and cave hyaena from several sites. The comparative measurements (in mm) have been extracted from Ehrenberg, 1940; Cardoso, 1993; García, 2003; Fourvel, 2012, and J.M.-M personal commun., 2015. Confidence ellipses with confidence interval (0.95) are figured.

FIGURE 12. 1, Graph representing total length of femur versus distal width of extant and cave hyaena from several sites. The comparative measurements have been extracted from Ehrenberg, 1940; Cardoso, 1993; Altuna and Mariezkurrena, 2000; Diedrich, 2011e. 2, Total length of tibia versus distal width of extant and cave hyaena from several sites. The comparative measurements have been extracted from Reynolds, 1902; Ehrenberg, 1940; Cardoso, 1993; Diedrich, 2011e; Fourvel, 2012. 3, Total length of third metatarsus versus smallest breadth (SD) of the diaphysis of extant and cave hyaena from several sites. The comparative measurements have been extracted from Ehrenberg, 1940; Cardoso, 1993; García, 2003; Fourvel, 2012. Confidence ellipses with confidence interval (0.95) are figured.