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Pleistocene cave hyenas in the Iberian Peninsula: New insights from Los Aprendices cave (Moncayo, Zaragoza)

Víctor Sauqué, Raquel Rabal-Garcés, Joan Madurell-Malaperia, Mario Gisbert, Samuel Zamora, Trinidad de Torres, José Eugenio Ortiz, and Gloria Cuenca-Bescós

Plain Language Abstract

Cave hyena (Crocuta spelaea) was a common carnivore in Europe, especially during Late Pleistocene, in spite of which there are very few examples of complete skeletons. In this paper, we present a new paleontological site called Los Aprendices, where the most complete skeleton in the Iberian Peninsula has been recovered. Besides, a taphonomic work has been done in order to understand the origin and preservation of these exceptional remains, suggesting a rapid burial after dead with scarce modification by other carnivores. The skeleton allows us to perform a biometric analysis and a study of the characteristics of cranial and appendicular skeleton, which result in the classification of cave hyenas as a different species and not as a subspecies of the extant spotted hyena (Crocuta crocuta). Additionally, a review of the Pleistocene record of genus Crocuta has been conducted, enabling us to establish one of the first records of Crocuta spelaea just in the recently discovered Los Aprendices cave.

Resumen en Español

Hienas de las cavernas del Pleistoceno en la Península Ibérica: nuevos conocimientos sobre la cueva de Los Aprendices (Moncayo, Zaragoza)

Se presenta un nuevo yacimiento paleontológico del Pleistoceno, Los Aprendices, ubicado en la parte noroeste de la Península Ibérica, en la zona del Moncayo (Zaragoza). La capa con restos fósiles ha sido datada por racemización de aminoácidos en 143,8 ± 38,9 miles de años (Pleistoceno Tardío más temprano o Pleistoceno Medio más tardío). Se han identificado cinco especies de mamíferos en la asociación: Crocuta spelaea (Goldfuss, 1823), Capra pyrenaica (Schinz, 1838), Lagomorpha indet., Arvicolidae indet. y Galemys pyrenaicus (Geoffroy, 1811). Los restos de C. spelaea corresponden a un esqueleto casi completo en semi-conexión anatómica. El espécimen de hiena representa el esqueleto más completo jamás recuperado en Iberia y uno de los restos más completos de Europa. Se ha comparado anatómica y biométricamente con las hienas de las cavernas europeas y las hienas manchadas actuales. Además, se ha realizado un estudio tafonómico para comprender el origen y la preservación de estos restos excepcionales. Los resultados sugieren un entierro rápido con pocas modificaciones por alteración producida presumiblemente por un carnívoro de tamaño medio. Se ha realizado una revisión del registro ibérico pleistoceno de Crocuta spp., que nos ha permitido establecer uno de los primeros registros de C. spelaea, en la recién descubierta cueva de Los Aprendices, y que nos ha mostrado también que la distribución geográfica más extensa de esta especie ocurrió durante el Pleistoceno Tardío (MIS4-2).

Palabras clave: Pleistoceno; Crocuta spelaea; Tafonomía; distribución; Península Ibérica

 

Traducción: Enrique Peñalver (Sociedad Española de Paleontología)

Résumé en Français

Les hyènes des cavernes pléistocènes de la péninsule Ibérique : nouvelles données de la grotte de Los Aprendices (Moncayo, Saragosse)

Un nouveau site paléontologique pléistocène, Los Aprendices, situé dans la partie nord-ouest de la péninsule Ibérique dans la région de Moncayo (Saragosse), est présenté. La couche avec les restes fossiles a été datée par racémisation des acides aminés à 143,8 ka ± 38,9 ka (base du Pléistocène récent ou toute fin du Pléistocène moyen). Cinq espèces de mammifères ont été identifiées dans l'assemblage : Crocuta spelaea (Goldfuss, 1823) Capra pyrenaica (Schinz, 1838), Lagomorpha indet., Arvicolidae indet., et Galemys pyrenaicus (Geoffroy, 1811). Les restes de C. spelaea comprennent un squelette sub-complet en semi-connexion anatomique. Le spécimen de hyène des cavernes représente le squelette le plus complet jamais trouvé dans la péninsule Ibérique et un des plus complets d'Europe. Des comparaisons anatomiques et biométriques ont été effectuées à la fois avec des hyènes des cavernes européennes et des hyènes tachetées actuelles. Une étude taphonomique a également été effectuée pour comprendre l'origine et la préservation de ces restes exceptionnels. Les résultats suggèrent un enfouissement rapide avec quelques modifications de charognage produites par un Carnivora de taille moyenne. Une synthèse du registre fossile pléistocène de Crocuta spp. dans la péninsule Ibérique a été effectuée, nous permettant de définir une des plus anciennes occurrences de C. spelaea dans la grotte récemment découverte de Los Aprendices, et montrant que la distribution géographique la plus vaste de cette espèce était au Pléistocène récent (SIM 4-2).

Mots-clés : Pleistocène ; Crocuta spelaea ; taphonomie ; distribution ; péninsule Ibérique

Translator: Antoine Souron

Deutsche Zusammenfassung

Pleistozäne Höhlenhyänen von der Iberischen Halbinsel: Neue Ergebnisse aus der Los Aprendices Höhle (Moncayo, Zaragoza)

Es wird eine neue pleistozäne paläontologische Fundstelle, gelegen im nordwestlichen Teil der Iberischen Halbinsel im Gebiet von Moncayo (Zaragoza), vorgestellt. Die fossilführende Schicht wurde mit Aminosäure-Racemisierung auf 143.8 ± 38.9 ka datiert (frühestes Spätpleistozän bis spätestes Mittelpleistozän). In der Assemblage wurden fünf Arten identifiziert: Crocuta spelaea (Goldfuss, 1823) Capra pyrenaica (Schinz, 1838), Lagomorpha indet, Arvicolidae indet und Galemys pyrenaicus (Geoffroy, 1811). Die Überreste von C. spelaea stellen ein nahezu komplettes Skelett in anatomischer Halbartikulation dar. Das Stück verkörpert das vollständigste Skelett, das jemals in Iberien gefunden wurde und eines der vollständigsten Überreste aus Europa. Es wurde anatomisch und biometrisch mit beiden europäischen Höhlenhyänen verglichen sowie mit den heutigen Tüpfelhyänen. Darüber hinaus wurde eine taphonomische Untersuchung durchgeführt, um den Ursprung und die Erhaltung dieser bemerkenswerten Überreste zu verstehen. Die Ergebnisse legen eine schnelle Einbettung nahe, mit wenigen Veränderungen durch Aasfresser, die vermutlich durch einen mittelgroßen Karnivoren verursacht wurden. Die pleistozänen iberischen Nachweise von Crocuta spp. wurden überprüft, was uns ermöglichte einen der frühesten Belege von C. spelaea in der kürzlich entdecken Los Aprendices Höhle etablieren. Außerdem wird herausgestellt, dass die größte geografische Verbreitung dieser Art während des späten Pleistozäns auftrat.

Schlüsselwörter: Pleistozän; Crocuta spelaea; Taphonomie; Verteilung; Iberische Halbinsel

Translator: Eva Gebauer

Arabic

Translator: Ashraf M.T. Elewa

 

 

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
Cervical V.   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
Mtc V R 1 1 1
Mtc II L 1 1 1
Mtc III L 1 1 1
Mtc IV L 1 1 1
Mtc V L 1 1 1
Femur R 1 10 10
Tibia R 1 2 2
Tibia L 1 1 1
Talus R 1 1 1
Calcaneus L 1 1 1
Calcaneus R 1 2 2
Cuboid L 1 1 1
Cuboid R 1 1 1
Pyramidal R 1 1 1
Scaphoid R 1 1 1
Scaphoid L 1 1 1
Cuneiform R 1 1 1
Cuneiform L 1 1 1
Mt II R 1 1 1
Mtt III R 1 1 1
Mtt IV R 1 1 1
Mtt V R 1 1 1
Mt II L 1 1 1
Mtt III L 1 1 1
Mtt IV L 1 1 1
Mtt V L 1 1 1
1st phalanx   12 16 16
2nd phalanx   8 9 9
3rd phalanx   4 4 4
Sesamoids   13 13 13
    107 194 194

 

TABLE 2. Overview of NR, number of remains; NISP, number of identified remains; MNE, minimum number of elements; MNI, minimum number of individuals and the distribution by age of the fossils from Los Aprendices.

          MNI by ages
Taxa NR NISP MNE MNI neonate adult
Capra pyrenaica 15 15 13 2 1 1
Crocuta spelaea 194 194 107 1   1
Unidentified (>4cm) 8          
Total 217 209 120 3 1 2

 

TABLE 3. Cave hyena remains (NISP) affected by carnivore bites in the skeleton of Crocuta spelaea from Los Aprendices.

  NISP Modified NISP % Modified
Axial skeleton 96 11 11.5
Zeugopod-stylopod 24 6 25
Autopod 71 3 4.2
Total 191 20 10.5

 

TABLE 4. Number of cave hyena remains (NISP) affected by different types of bites in the skeleton of Crocuta spelaea from Los Aprendices.

  "Groove" "Pit" "Puncture" Crenulated edge Impact point "Furrowing" "Scoping out"
Cervical V.           2  
Thoracic V.           1  
Lumbar V.   3   2   1  
Indet. V.   1 1 1 1 2  
Scapula     1     1  
Ribs   1 1 2   2  
Humerus 1 1 1   1   1
Ulna   1 1   1 2  
Femur 1         1  
Tibia 1 1 2     1 1
Tarsal 1         1  
Phalanx           2  
NISP 4 8 7 5 3 16 2

 

TABLE 5. Measurements of the “pits”, “punctures” and grooves found in the skeleton of Crocuta spelaea from Los Aprendices. Abbreviations: min, minimum; max, maximum; σ, standard deviation; n, sample size; L, length; W, width.

  Pits in
cancellous bone
Punctures in
cancellous bone
"Pits" & "punctures" in cancellous bone Grooves in cancellous bone
  Width Length L/W Width Length L/W Width Length L/W Width Length
Mean 2.38 3.21 1.35 3.61 4.49 1.2 2.91 3.77 1.31 1.6 5.7
σ 0.76 1.27 1.67 1.32 1.46 1.1 1.14 1.46 0.26 0.57 1.48
Min. 1.49 1.59 1.07 2.19 3.33 1.5 1.49 1.59 0.92 1.01 3.66
Max. 3.35 5.16 1.54 6.4 7.42 1.2 6.4 7.42 1.73 2.33 7.55
n 9 9   7 7   17 16 16 8 8

 

TABLE 6. Regression equations used for estimating the body mass of Crocuta spelaea and estimates of size obtained with them for the living hyenas modified of Pamlqvit et al., 2011. Deviations between estimated and actual masses for extant species [(mass estimate)/mass*100] are shown between brackets. Abbreviations: X, mean; min, minimum; max: maximum; BM, body mass; SKL, skull length; Lm1, lower carnassial length; HL, humerus length; FL, femur length.

References Equation Los Aprendices Crocuta spelaea   Crocuta crocuta
(55 kg)
  Pachycrocuta brevirostris
N X min max   N    
Van Valkenburgh, 1990 log10 (BM)= 3.13log10 (SKL) - 5.59 145.6 22 140 107.2 173.1   72 63 (14.5%)   176
  log10 (BM)= 2.72log10 (Lm1) - 2.03 135.6 177 112.6 73 165     62 (12.7%)   92
Anyonge, 1993 log10 (BM)= 2.93log10 (HL) - 5.11   19 75.3 58.9 93.5   28 54 (-1.9%)   115
  log10 (BM)= 2.92log10 (FL) - 5.27 67.5 15 69.7 57.1 86.6     52 (-5.5%)   86
Christiansen, 1999 log10 (BM)= 2.88log10 (HL) - 5.15   19 65.7 51.65 81.3   30 48 (-12.7%)   89
  log10 (BM)= 2.86log10 (FL) - 5.15 63.6 15 65.6 54 81.2   199 49 (-10.9%)   80
Andersson, 2004 log10 (BM)= 2.55log10 (TC) - 0.6               62 (12.7%)   110
Average   103   88.15         56 (1.8%)   108
 

sauqueVíctor Sauqué. Grupo Aragosaurus-IUCA, Dpto. Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009, Zaragoza, Spain; This email address is being protected from spambots. You need JavaScript enabled to view it.
Laboratorio de Zooarqueología y Tafonomía de Zonas Áridas (LaZTA), IDACOR, CONICET/FFyH UNC. Av. H. Yrigoyen 174, 5000, Córdoba, Argentina

Víctor Sauqué obtained his Geology Degree in 2008, being valedictorian of his class, a Master in Geological Science in 2009, a Master in Science Teaching in 2015 and his PhD in Geological Sciences in 2015 with a thesis titled "Taphonomy, systematics and paleoenvironmental approach of Upper Pleistocene macromammals of Moncayo (Zaragoza)". Recently he directed excavations campaigns in Los Rincones and Los Aprendices (Zaragoza) and in the last two years he led a project at high altitude in the Pyrenees, over 2000 m.a.s.l. In these campaign, the team discovered several new sites with interesting remains which are under study at this very moment. He is Research Collaborator at the Natural Science Museum Zaragoza. He is currently a researcher at the Universidad Nacional de Córdoba, Argentina. Apart from his scientific resume, in the past years he has been involved in education programs to promote science amongst young students at school in the program "Little palaeontologist" directed by Federico Ozanam Foundation.

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rabalRaquel Rabal-Garcés. Grupo Aragosaurus-IUCA, Dpto. Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009, Zaragoza, Spain; This email address is being protected from spambots. You need JavaScript enabled to view it.

Dra. Raquel Rabal Garcés has a degree in geology from the University of Zaragoza (2006), Master's in Introduction to Research in Geology (2008) and PhD in Geology from the University of Zaragoza (2013). In her thesis, entitled "Estudio paleontológico de Ursus spelaeus Rosenmüller, 1794 del Pleistoceno Superior de Coro Tracito (Tella, Huesca, España)" conducted a detailed study of the taphonomy, systematic paleontology and paleoecology of fossil remains of Ursus spelaeus in order to characterize the population.

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madurellJoan Madurell-Malaperia. Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c\de les columnes sn, 08193, Cerdanyola del Vallès, Barcelona, Spain; This email address is being protected from spambots. You need JavaScript enabled to view it.

Joan Madurell-Malapeira, currently works as a postdoctoral researcher at the Catalan Institute of Palaeontology. He obtained his PhD in Geological Sciences in December of 2010 with a thesis entitled "The Villafranchian carnivores of the North-Eastern Iberian Peninsula and the Early-Middle Pleistocene Transition", under the direction of Prof. Salvador Moyà-Solà and Prof. Bienvenido Martínez-Navarro. From 2007 onwards, he has published more than thirty papers in scientific journals including, Quaternary Science Reviews, Journal of Human Evolution, Journal of Vertebrate Paleontology and Molecular Ecology. These papers are mostly devoted in Carnivore taxonomy and paleobiology and their relationships of competence with the first European hominins. This research is also focused in the consequences of the great climatic shifts associated to the 'Mid-Pleistocene Revolution' in the large mammal assemblages and
specially in the competition between the large felids, hyenas and the first European hominins. The last three years, he started a new project devoted in the endocranial morphology of the Miocene to Pleistocene large carnivores, basically felids and hyenids, in order to make inferences in the degree of sociability of these carnivores. Furthermore has a lot of experience in fieldwork, having taken part in many excavations at several fossil localities from the Vallès- Penedès Basin, Banyoles-Besalú Basin and Guadix-Baza Basin (Spain). His capacity of leadership is evidenced by the fact that, among others, he was director of the paleontological interventions of Vallparadís Estació (Terrassa, Catalonia, Spain) from 2005 to 2008 and director of the paleontological interventions in the Incarcal complex from 2004 to the present.

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gisbertMario Gisbert. Centro de espeleología de Aragón (CEA); This email address is being protected from spambots. You need JavaScript enabled to view it.

Mario Gisbert León is a researcher who collaborates with the Department of Paleontology of the University of Zaragoza. He is a member of the Spanish Society of Speleology and Study of the Karst, the Center of Speleology of Aragon and the Grackma, Glacial, Criokarst and Environment Association. He has supervised numerous research projects, topography and underground exploration in various kartic zones of Europe and South America. He is a specialist in techniques of vertical progression and underground rescue.

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zamoraSamuel Zamora. Instituto Geológico y Minero de España, Calle Manuel Lasala, 44, 9ºB, 50006, Zaragoza, Spain; This email address is being protected from spambots. You need JavaScript enabled to view it.

Samuel Zamora studied geology at the University of Zaragoza, Spain, graduating in 2004. He completed his PhD in 2009 at the same university, studying Cambrian echinoderms from North Spain, including faunas from the Moncayo Natural Park. During 2010–2012, he spent two years working as a postdoc at the Natural History Museum, London, and in 2013, he was awarded a Postdoctoral Fellowship at the Smithsonian Institution National Museum of Natural History, and he was based in Washington D.C., USA for one year. He currently works at the Spanish Geological Survey funded by a Ramón y Cajal grant and he is Research Collaborator at the Smithsonian Institution.

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torreTrinidad de Torres. Laboratory of Biomolecular Stratigraphy, Madrid School of Mines; This email address is being protected from spambots. You need JavaScript enabled to view it.

Trinidad de Torres is a Full Professor at the School of Mines in Madrid. In this institution he leads a research group in amino acid racemization dating method. He obtained his pHD in the study of bears "Ursids of the Pleistocene-Holocene of the Iberian Peninsula".

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ortizJosé Eugenio Ortiz. Laboratory of Biomolecular Stratigraphy, Madrid School of Mines This email address is being protected from spambots. You need JavaScript enabled to view it.

Biomolecular Stratigraphy Laboratory. Madrid School of Mines and Energy. Universidad Politécnica de Madrid. C/Ríos Rosas 21, Madrid 28002, Spain José E. Ortiz is a PhD Mining Engineer and a professor of Geology. He is the secretary of the Geological Society of Spain. His research is related to stratigraphy, organic geochemistry and palaeoclimatology. He is interested in the amino acid racemization of ostracodes, molluscs and vertebrates.

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cuencaGloria Cuenca-Bescós. Grupo Aragosaurus-IUCA, Dpto. Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009, Zaragoza, Spain; This email address is being protected from spambots. You need JavaScript enabled to view it.

Gloria Cuenca-Bescós is a Senior Lecturer in Geology at the Department of Earth Sciences at the University of Zaragoza in Spain. She focus her research in the small mammals, their fossils from the time of the Dinosaurs to the Late Pleistocene and up to the present day. She is involved in the excavations and scientific studies of several sites in Spain, being interested in the paleobiology and biochronology of the fossil vertebrate assemblages in the stratigraphic sequences and in comparison with other sites from Eurasia. Also she is involved in the preservation of the paleontological record troughout her contribution to the Museum of Natural Sciences (MusNacUnizar) and Environmental Sciences Institute (IUCA) both of the University of Zaragoza.

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FIGURE 1. 1, Metapodials of Crocuta spelaea in anatomical semi-connection. 2, Cranium of Crocuta spelaea. 3, Filtering the excavation grid. 4, Excavation of various Crocuta spelaea remains from the hyena level.

figure1 

FIGURE 2. 1, Geographical situation of Los Aprendices Cave. 2, Panoramic view of the western slopes of the Peñas del Cabo. 3, Mouth of Los Aprendices Cave. 4, Main gallery of the cave.

figure2 

FIGURE 3. 1, Elevation view of Los Aprendices Cave. 2, Plan view of Los Aprendices Cave. 3, Detailed elevation view of the hyena gallery showing the excavation grid. 4, Detailed plan view of the hyena gallery showing the excavation grid as well as how the squares that were excavated were coloured. 5, Sediment cone that blocks off the former entrance to the hyena gallery. Within it one can see the hyena level, from which the skeleton of Crocuta spelaea was recovered.

 figure3

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

figure4 

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

figure5 

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). 

 figure6

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).

figure7 

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.

figure8 

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.

 figure9

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.

 figure10

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.

 figure11

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.

figure12 

FIGURE 13. 1, Analysis of the breakage of the long bones from the skeleton of Crocuta spelaea from Los Aprendices, showing the abundance of each type of fracture according to the criteria analysed: angle, delineation and edge of the breaks. 2, Analysis of the breakage of the diaphyses in terms of circumference (C1, C2, C3) and length (L1, L2, L3, L4) for the skeleton of Crocuta spelaea from Los Aprendices.

figure13 

FIGURE 14. Photographs of the various types of marks (indicated by white arrows) present on the skeleton of Crocuta spelaea from Los Aprendices. 1, detailed image of the proximal epiphysis of the right humerus (MPZ 2014/672). 2, humerus in cranial view (MPZ 2014/672). 3, humerus in lateral view (MPZ 2014/672). 4, right ulna (MPZ 2014/676) showing “furrowing” on the proximal epiphysis, as well as a “puncture”. 5, left ulna (MPZ 2014/610) showing “furrowing” on the proximal epiphysis. 6, right tibia (MPZ 2014/681) showing “furrowing” on the proximal epiphysis. 7-8, lateral and dorsal views of a left calcaneus (MPZ 2014/677) showing “furrowing” and grooves on the calcaneal tuberosity. 9, rib (MPZ 2014/591) showing a “puncture”. 10, lumbar vertebra (MPZ 2014/582) in caudal view showing “furrowing” on the vertebral body. 11, lumbar vertebra (MPZ 2014/582) in ventral view showing three pits, corresponding to the incisors of a medium-sized carnivore. 12-13, lateral and caudal views of a dorsal vertebra (MPZ 2014/674) showing how the vertebral body has been torn off by a bite.

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FIGURE 15. Palaeogeographic and diachronic distribution of the Crocuta in the Iberian Peninsula. The numbers refer to the sites listed in Appendix 7.

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FIGURE 16. Graph representing total length of femur versus total length of third metatarsal in the same individual of extant Crocuta crocuta and the three unique complete skeletons of Crocuta spelaea from three sites Koněprusy Cave (Diedrich, 2012a); Gargas (Cardoso, 1993) and Los Aprendices this work. Comparative measurements (in mm) have been extracted from Ehrenberg, 1940; Cardoso, 1993; and J.M.-M personal commun., 2015. Confidence ellipse with confidence interval (0.95) is figured.

 figure16

FIGURE 17. Means (square) and standard deviations (DS) of the values for the width (1) and the length (2) of pits on the epiphyses of long bones for various modern-day carnivores, together with the data from Los Aprendices (arrows). Key to the data: “Selvaggio and Wilder, 2001; ‘ Domínguez-Rodrigo and Piqueras, 2003; ^ Delaney-Rivera et al., 2009; * Saladié, 2009.

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FIGURE 18. Reconstruction of Crocuta spelaea in the vicinity of Los Aprendices Cave, feeding on the carcass of a specimen of Capra pyrenaica (illustration by Gianfranco Mensi).

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APPENDIX 1.

Abundance, racemization relations of aspartic acid (Asp), glutamic acid (Glu), serine (Ser) and age of the samples from Los Aprendices. D: dextro; L :levo; ka: kilo annum; mg: milligram; pmol: picomol. All appendix files are available at the website as zipped spreadsheet files.

APPENDIX 2.

Cranial measurements (in mm) of cave hyena from the Iberian Peninsula compared to other European individuals and extant individuals. Abbreviations: L, Length; H, height; W, width; Ap, Los Aprendices; Ga, Gabasa; Fo, Fontainhas; Ga, Gargas; Ch, Châtillon Saint Jean; Ja, Jaurens; Bi, Binagady; Jav, Javorka Cave; Sl, Sloup Cave; Ka, kazanka; Mo, Mosbach; LV, Lunel-Viel; TDW, Atapuerca Trinchera Dolina, extant Crocuta crocuta ; A, Blasco and Montes, 1997; b, Cardoso, 1993; c, J.M.-M personal commun., 2015; d, Baryshnikov, 2014; e, Bonifay, 1971; f, García, 2003; g, Baryshnikov and Tsoukala, 2010. Note:? denotes uncertainity in the measure due to a fracture in the bone. All appendix files are available at the website as zipped spreadsheet files.

APPENDIX 3.

Upper dentition measurements (in mm) of Los Aprendices Crocuta spelaea compared to other fossil hyenas and extant Crocuta crocuta. Abbreviations: X: mean; min; minimum; max: maximum; DMD, Mesiodistal diameter; DVL, vestibulo-lingual diameter; Ko, Kobaederra;Fo, Fontainhas; Ag, Aguilón; Ka, Kazanka; Mo, Mosbach; a, Altuna and Mariezkurrena , 2000; b, Castaños, 1987; c, Blasco and Montes, 1997; d, Iñigo et al., 1998; e, Cardoso, 1993; f, J.M.-M and personal commun., 2015; g, García, 2001; h, Bonifay, 1971; I, Kurtén and Poulianos, 1981; j, Kurtén, 1962; k, Schütt, 1971; l, Baryshnikov, 2014; m, Turner, 1984; n, Hollister, 1918. All appendix files are available at the website as zipped spreadsheet files.

APPENDIX 4.

Mandible measurements (in mm) of Los Aprendices Crocuta spelaea compared to other fossil hyenas and extant Crocuta crocuta. Abbreviations: X: mean; min; minimum; max: maximum; Hramus, mandibular ramus height; Hdiast, height of ramus below the diastema; Hm1, height of ramus below m1; Hp4, height of ramus below p4; H condyle md, height of mandibular condyle; W condyle md, width of the mandibular condyle; Ap, Los Aprendices; Ga, Gabasa; Va, Valdegoba; S.Ab, Sima de Abraham; Ca, Caldeirao; Gar, Gargas; Eh, Ehringsdorf; Te, Teixoneres; TR, Terrasses des Canyars; Ka, Kazanka; a, Blasco and Montes, 1997; b, García, 2001; c, Altuna and Mariezkurrena, 2000; d, Martínez-Sánchez et al., 2012 e, Davis, 2002; f, Cardoso, 1993; g, Rosell et al., 2012; h, Daura et al., 2013; i, Baryshnikov, 2014; j, Bonifay, 1971; k, J.M.-M personal commun., 2015. All appendix files are available at the website as zipped spreadsheet files.

APPENDIX 5.

Lower dentition (in mm) of Los Aprendices Crocuta spelaea compared to other fossil hyenas and extant Crocuta crocuta. Abbreviations: X: mean; min; minimum; max: maximum; DMD, Mesiodistal diameter; DVL, vestibulo-lingual diameter; Ap, Los Aprendices: L.K, Labeko Koba; S.I, Sima I; Bo, Bolinkoba; Ga, Gabasa; Fo, Fontainhas; Va, Valdegoba; C.B, Cueva del Búho; To, El Toll; Ol, Olopte B; C.O, Cova de l’Or; Mo, Mollet; Ata, Atapuerca; LV, Lunel-Viel; Pe, Petralona; Mos, Mosbach; Ka, Kazanka; a, Altuna and Mariezkurrena , 2000; b, Castaños, 1987; c, Blasco and Montes, 1997; d, Cardoso, 1993; e, García, 2001; f, Iñigo et al., 1998; g, J.M.-M personal commun., 2015; h, Bonifay, 1971; I, Kurtén and Poulianos, 1981; j, Schütt, 1971; l, Turner, 1984; m, Hollister, 1918. All appendix files are available at the website as zipped spreadsheet files.

APPENDIX 6.

Postcranial measurements (in mm) of Los Aprendices Crocuta spelaea compared to other fossil hyenas and extant Crocuta crocuta. Abbreviations: X: mean; min; minimum; max: maximum, Bd: Breadth of the distal end; Bp: Breadth of proximal end; BPC, Proximal articular surface breadth; DPA, Olecranic maximum depth; GB: Greatest breadth; GL: Greatest length; SD: smallest breadth of the diaphysis; Mtc: metacarpus; Mtt: metatarsus; 1st Pha: first phalanx; 2nd Pha: second phalanx; Ap, Los Aprendices; L.K, Labeko Koba; S.I, Sima I; Ga, Gabasa; To, El Toll; TD 8, Atapuerca Trinchera Dolina 8; LV, Lunel-Viel; A, Altuna and Mariezkurrena , 2000; b, Blasco and Montes, 1997; c, Castaños, 1987; d, García, 2001; e, J.M.-M personal communication., 2015; f, Cardoso, 1993; g, Bonifay, 1971; h, Ehrenberg, 1940. Note: ? denotes uncertainity in the measure due to a fracture in the bone. All appendix files are available at the website as zipped spreadsheet files.

APPENDIX 7.

Palaeontological and archaeological sites of the Iberian Peninsula with hyena remains. The numbers refer to the sites of Figure 1. Abbreviations: Context: A, archaeological; P, palaeontological. Chronology: P, Pleistocene; MP, Middle Pleistocene; ELP, Early Late Pleistocene; LP, Late Pleistocene; EH, Early Holocene; MPal, Middle Palaeolithic; EUPal, Early Upper Palaeolithic; UPal, Upper Palaeolithic; Ms, Mousterian; Chat, Chatelperronian; Per, Perigordian; Au, Aurignacian; Gr, Gravettian; So, Solutrean; Mg, Magdalenian; Az, Azilian. All appendix files are available at the website as zipped spreadsheet files.