Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe

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Geobios 41 (2008) 677–687 http://france.elsevier.com/direct/GEOBIO/

Original article

Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe Taxonomie et mode`les e´volutifs des Hyaenidae fossiles d’Europe Alan Turner a,*, Mauricio Antón b, Lars Werdelin c b

a School of Biological and Earth Sciences, Liverpool John Moores University, L3 3AF Liverpool, UK Departamento de Paleobiología, Museo Nacional de Ciencias Naturales, C/José Gutiérrez Abascal 2, 28006 Madrid, Spain c Department of Palaeozoology, Swedish Museum of Natural History, Box 50007, 104 05 Stockholm, Sweden

Received 25 October 2007; accepted 9 January 2008

Abstract We review the larger pattern of appearance of the Hyaenidae in Europe and outline their part in the turnover of the guild of larger Carnivora that occurs across the Miocene–Pliocene boundary. The earliest record of the family is in MN4, although the patchy nature of the earliest records makes it difficult to be certain about the continent of origin. There is a clear pattern of morphological evolution over that long timespan, from the earliest viverrid- and herpestid-like forms through dog-like and more cursorial taxa to the larger, bone-crunching animals of the later Miocene and the Pliocene–Pleistocene epochs. Miocene dog-like hyaenas may indicate that social hunting had emerged by that time, while the appearance of larger species means that hyaena-accumulated bone assemblages may potentially occur in any late Miocene to Pleistocene locality. # 2008 Elsevier Masson SAS. All rights reserved. Résumé Nous révisons le modèle d’apparition des Hyaenidae en Europe et nous soulignons leur rôle dans le renouvellement des communautés de grands carnivores au cours du Miocène et du Pliocène. Le premier enregistrement de cette famille est situé dans la MN4, même si la rareté des premiers enregistrements ne permet pas d’établir avec certitude leur continent d’origine. Il existe un modèle clair d’évolution morphologique pendant cette période, des premières formes apparentées aux viverridés et aux herpestidés en passant par des formes de type dog-like et des taxons plus cursoriaux, jusqu’aux grands animaux broyeurs d’os du Miocène terminal et du Plio-Pléistocène. Les hyènes miocènes qualifiées de dog-like peuvent indiquer que la chasse sociale, en groupe, ait émergé à cette époque, alors que l’apparition des espèces de grande taille signifie que les hyènes accumulatrices de vestiges osseux, ont pu potentiellement exister dans les localités du Miocène terminal au Pliocène. # 2008 Elsevier Masson SAS. All rights reserved. Keywords: European Hyaenidae; Guild evolution; Social hunting; Bone accumulators Mots clés : Hyaenidae d’Europe ; Évolution de guilde ; Chasse sociale ; Accumulateurs d’os

1. Introduction The Hyaenidae are a taxonomically diverse family of Neogene Carnivora. The three larger living species contribute significantly to the structure of their predatory guild and the nature of their food debris may offer much insight into the ecosystem within which they operate (Kruuk, 1972; Mills, 1990). Many European localities of Miocene–Pleistocene age

* Corresponding author. E-mail address: [email protected] (A. Turner).

contain hyaenas and a significant number of Middle and, particularly, Upper Pleistocene cave assemblages consist largely of the remains of hyaenas and their prey (Kurtén, 1968). The structural composition of the family, seen in terms of size and morphology of its constituent species, has changed over time across the entire Afro-Eurasian extent of its distribution (Werdelin and Turner, 1996a, 1996b). Hyaenas are of significant palaeontological interest, but any investigations must operate within a clearly understood taxonomic framework. The detailed review by Werdelin and Solounias (1991) has gone a long way to providing such a framework, while a general revision of African Pliocene and

0016-6995/$ – see front matter # 2008 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2008.01.001

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Pleistocene taxa has been undertaken (Turner, 1990; Werdelin and Turner, 1996a) and a detailed treatment of the giant hyaena, Pachycrocuta brevirostris (Aymard) recently published (Turner and Antón, 1996). Werdelin and Solounias (1996) have reviewed the evolutionary history of the Miocene Hyaenidae of Europe and western Asia. Here, we provide a taxonomicallybased overview of the European members of the family to parallel that previously published on the African taxa (Werdelin and Turner, 1996a) and consider some of the evolutionary patterns seen in the family, in the hope that this will set the study of Plio-Pleistocene hyaenas within a larger palaeoecological context. We confine our attention to taxa found in Europe proper, although we omit taxa only found in Moldova and Ukraine and give some mention to wider distributions of European hyaenas. 2. Hyaenid taxa Over 60 probable species of hyaena have so far been identified (Werdelin and Solounias, 1991, 1996; Werdelin and Turner, 1996a). Fig. 1 is a cladogram of the family based on these and later studies. We exclude the genera Percrocuta and Dinocrocuta from the Hyaenidae and place them in a separate family, the Percrocutidae (Werdelin and Solounias, 1991), although one genus tentatively excluded by Werdelin and Solounias, Allohyaena, is now restored to the Hyaenidae (Werdelin and Kurtén, 1999). Like the former authors, we have employed the convention of indicating generic names in quotation marks for species lacking sufficient characters for unambiguous allocation to monophyletic genera. Our treatment includes allocation of the included species to one of six categories of adaptive types – in effect grades and referred to here as ‘‘Ecomorph Groups’’ – devised by Werdelin and Solounias (1996). Briefly, the six categories are:      

civet-like insectivore/omnivore; mongoose-like insectivore/omnivore; jackal- and wolf-like meat and bone eaters; cursorial meat and bone eaters; transitional bone crackers; fully-developed bone crackers.

This system has proved useful in our previous investigation of the African Hyaenidae (Werdelin and Turner, 1996a), since it provides a convenient overview of the structural evolution of the family, although probably necessitating some revision to the separation of the first two groups, which both retained generalised climbing abilities. Some of these morphological types are shown in Fig. 2 and 3. For individual species, we offer a list of localities where these are few in number although with Pliocene and later taxa, a detailed account quickly becomes unmanageably large and such lists serve more as a general guide. Genus Protictitherium Kretzoi This genus was created by Kretzoi (1938, 1984), one of the first to recognise the true complexity of the evolutionary history

of the Hyaenidae. Werdelin and Solounias (1991, 1996) showed that the genus includes the earliest and most primitive members of the family, coming from deposits allocated to MN4 (although surviving to latest Miocene times) and placed them within Ecomorph Group 1 of their scheme. They share a generalised civet-like dentition, with a complete complement of premolar and molar teeth and a postcranial skeleton that shows essentially terrestrial features and retractile claws with asymmetrical second phalanges (Semenov, 1989), indicating that climbing was undoubtedly possible. ‘Protictitherium’ gaillardi (Forsyth Major) Astaracian-Vallesian (MN4–10). Known from Vieux-Collognes, Bézian, Castelnau-d’Arbieu, La Grive Saint-Alban Simorre, Langenau 1 and Pontlevoy in France, Arroyo del Val IV Can Llobateres, Can Llobateres I, Can Ponsic, Can Ponsic I, Castell de Barbera, Hostalets de Pierola, Santiga, Terrassa and Paracuellos del Jarama in Spain. This species and P. crassum (below) are separated largely by size, with ‘Protictitherium’ gaillardi as the smaller of the two (Werdelin and Solounias, 1991). Protictitherium crassum (Depe´ret) Astaracian-?Turolian (MN5–12). Known from Castelnaud’Arbieu, La Grive, Saint-Alban and Montredon in France, Dorn Dürkheim and Eppelsheim in Germany, Xirochori 1 and Dytiko in Greece, Kalfa in Moldova, Can Llobateres I and II, Los Valles de Fuentedueña, Hostalets de Pierola Inferior, Hostalets de Pierola Superior, La Retama, Santiga and Cerro Batallones in Spain as well as localities in Turkey and the Crimea. This species and P. llopisi were previously separated at the subspecific level by Crusafont Pairó and Petter (1969), but Werdelin and Solounias (1991) argued for specific-level differences between the two. Protictitherium crassum is illustrated here in Fig. 2. ‘Protictitherium’ llopisi (Crusafont Pairo´ and Petter) Vallesian-Turolian (MN11–12). Known from Can Bayona in Spain. ‘Protictitherium’ sumegense Kretzoi Turolian (MN11). Known from Sümeg in Hungary. Werdelin and Solounias (1991) pointed out that this species has never been figured since its proposal by Kretzoi (1984) and is in any event, poorly characterised. The material, consisting of a few isolated teeth, is not sufficient to distinguish this species from others in the genus and we therefore consider it a nomen dubium. Protoctitherium csakvarense Kretzoi Turolian (MN11). Known from Csákvár in Hungary. A poorly characterized species only doubtfully distinct from better known Protictitherium species, such as Protoctitherium crassum (see Kretzoi, 1951).

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Fig. 1. Our current view of the phylogeny of the Hyaenidae. Species discussed in this paper are denoted by asterisks. The type species of each genus has the name spelled out in full. Notre interprétation actuelle de la phylogénie des Hyaenidae. Les espèces discutées sont indiquées par un astérisque. L’espèce typique de chaque genre est épelée complètement.

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Fig. 2. Some Miocene hyaenas. From left to right: Adcrocuta eximia, Hyaenotherium wongii, Ictitherium viverrinum, Protictitherium crassum and Plioviverrops orbignyi. Scale bar is 25 cm. Quelques hyènes miocènes. De gauche à droite : Adcrocuta eximia, Hyaenotherium wongii, Ictitherium viverrinum, Protictitherium crassum et Plioviverrops orbignyi. Échelle : 25 cm.

Genus Plioviverrops Kretzoi Members of this genus are the sole constituents of Ecomorph Group 2, with a broadly mongoose-like insectivore/omnivore morphology that includes a reduction in the sectorial part of the dentition, retention of moderate claw retraction and a postcranial skeleton generally more adapted to a terrestrial mode of life. They are the second oldest group of hyaenas, ranging in time from MN5 to 13 and possibly 14. ‘Plioviverrops’ collectus (De Bonis) This species is known from the MN2 locality Laugnac in France. De Bonis (1994) has recently transferred this material, previously referred to the then new species Herpestides

collectus De Bonis to the hyaenid genus Plioviverrops. If correct, this would make Plioviverrops collectus the oldest known true hyaenid. Unfortunately, de Bonis’ text and accompanying illustrations provide no answer to why the material was transferred to Plioviverrops. He argues convincingly that the material should indeed be removed from Herpestides, a viverrid, but then states that the characters typifying the specimens ally them with Plioviverrops without discussing what these characters might be or providing any distinction between primitive and derived features. The characters used to distinguish the Laugnac material from Herpestides are mainly features that are derived in the latter genus and indicate no special relationship to Plioviverrops or to the Hyaenidae. Thus, the generic and familial identity of this

Fig. 3. Some Pliocene and Pleistocene hyaenas. From left to right: Crocuta crocuta, Pliocrocuta perrieri (foreground), Pachycrocuta brevirostris (background) and Chasmaporthetes lunensis. Scale bar is 25 cm. Quelques hyènes pliocènes et pléistocènes. De gauche à droite : Crocuta crocuta, Pliocrocuta perrieri (premier plan), Pachycrocuta brevirostris (arrière plan) et Chasmaporthetes lunensis. Échelle : 25 cm.

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material must remain uncertain and cannot be used to demonstrate the presence of Hyaenidae in MN2. Plioviverrops gervaisi De Beaumont and Mein Orleanian (Upper Burdigalian) (MN4–5). Known from Vieux-Collognes in France and perhaps from Catalayud in Spain. Werdelin and Solounias (1991) raise the question of whether this species is truly distinct from Plioviverrops gaudryi. Both species are known only from sparse dental remains. Plioviverrops gaudryi De Beaumont and Mein Astaracian (MN7–8). Known from La Grive Saint-Alban in France. Plioviverrops orbignyi (Gaudry and Lartet) Vallesian-Turolian (MN10–12). Known from Pikermi, Ravin des Zouaves 5, Samos main Bone Beds, Prochoma, Vathylakkos and Perivolaki in Greece and Los Aljezares in Spain. This species, illustrated in Fig. 2, is one of the youngest and has been suggested by Thenius (1966) as a plausible ancestor of the aardwolf, Proteles cristatus (Sparrman), a suggestion with which Werdelin and Solounias (1991) concurred in the absence of evidence to the contrary. However, recent molecular work by Koepfli et al. (2006) suggests that the divergence time of Proteles from the other extant hyenas is too late (at c. 0.6 Ma) for it to be derived from Plioviverrops, as the divergence of this genus from other Hyaenidae goes back to MN4 (18–17 Ma). Plioviverrops guerini (Villalta Comella and Crusafont Pairo´) Turolian (MN11–12). Known from Crevillente-2, Piera, Puente Minero, Vivero de Pinos, Cerro de la Garita, Concud and Los Mansuetos in Spain. Again represented by sparse material. ‘Plioviverrops’ faventinus Torre Ruscinian (MN13–14). Known from Brisighella in Italy and La Gloria 4 in Spain, this is the species in which the Ecomorph 2 trend towards an omnivore/insectivore morphology culminates. The reported occurrence of this species at La Gloria 4 (Alcala et al., 1992) extends the time-range of the genus into the Pliocene and Alcala (1994) has discussed the material at length. This is one of the few carnivore taxa whose temporal range crosses the Miocene–Pliocene boundary (Werdelin and Turner, 1996b). Genus Thalassictis Gervais This genus, together with the genera Ictitherium and Hyaenictitherium falls within Ecomorph Group 3. Members of this group have a generalised canid-like dentition, and have been likened to jackal- and wolf-like bone and meat eaters (Werdelin and Solounias, 1996; Werdelin and Turner, 1996a). Thalassictis is the earliest-known member of the group, with its first occurrence in MN7–8 deposits and shows a characteristic

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reduction of the posterior molars and an increase in body size. Werdelin and Solounias (1991) have discussed the exceedingly complex history of the nomenclature of this genus and its various constituent species in detail. ‘Thalassictis’ certa (Forsyth Major) Astaracian (MN7–8). Known from La Grive Saint-Alban in France. ‘Thalassictis’ montadai (Villalta Comella and Crusafont Pairo´) Upper Astracian-lower Vallesian (MN7–9). Known from Ballestar, Can Barra, Can Mata 2, Can Ponsic I and Hostalets de Pierola Superior and Inferior in Spain and also from Yeni Eskihisar in Turkey. Thalassictis robusta Gervais Vallesian-Turolian (MN9–11). Known from Höwenegg and Dorn Dürkheim in Germany. Identification of the German material was made by De Beaumont (1986). Werdelin and Solounias (1991) expressed strong reservations, but accepted the allocation in the absence of better data. Genus Ictitherium Roth and Wagner This long-established genus, the second of the Ecomorph Group 3 genera, appears in MN9 deposits. Werdelin and Solounias (1996) suggest that it may either derive from previous Group 3 taxa or independently from Group 1. Members of the genus are characterised by a lesser reduction of the posterior molars and less sectorial carnassials than Thalassictis. Ictitherium viverrinum (Roth and Wagner) Vallesian-Turolian (MN9–13). Known from Montredon in France, Vösendorf in Austria, Titov Veles in the Former Yugoslavian Republic of Macedonia (FYROM), Pikermi, Ravin des Zouaves 5, Samos Main Bone Beds and White Sands and Vathylakkos in Greece. Also known from Belka and Grebeniki in the Ukraine and Upper Maragheh in Iran and across to Lufeng in China. This, the type species of the genus Ictitherium, is another taxon with a long and complex taxonomic history involving confusion with material now referred to Thalassictis robusta, as reviewed by Werdelin and Solounias (1991). Comments on the postcrania were made by both Gaudry (1862–1867) and Pilgrim (1931) with the former suggesting proportional similarities to a civet, while Pilgrim pointed to a relatively shortened hindlimb (see Fig. 2). Ictitherium pannonicum Kretzoi Turolian (MN12–13). Known from Polgárdi in Hungary and possibly Valdecebro in Spain. Werdelin and Solounias (1991) give a detailed discussion to the relationships of this taxon, first established by Kretzoi (1952) and with its validity more recently supported by Semenov (1985).

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‘Ictitherium’ adroveri (Crusafont Pairo´ and Petter) Turolian (MN12–13). Known from Arquillo and Los Mansuetos in Spain. Werdelin and Solounias (1991) pointed to the doubtful validity of this species, based as it was on two isolated teeth, which could not even be referred to the same taxon with any certainty. They, therefore, suggest that the taxon be considered a nomen dubium. More recently, Alcala (1994) has discussed the matter and concluded that the taxon is a synonym of Hyaenictitherium wongii. Genus Hyaenictitherium Kretzoi The genus Hyaenictitherium, the last of the Group 3 members, was erected by Kretzoi (1938) in his effort to sort out the confusion produced by earlier taxonomic schemes. It is essentially an Asian taxon and specimens assigned to the genus have been recovered from Chinese and Siwalik deposits of broadly Turolian age (Werdelin and Solounias, 1991). The sole European species is Hyaenictitherium wongii (see below), which has a wide distribution in the Upper Miocene of China. Hyaenictitherium wongii (Zdansky) Vallesian-Turolian (MN9–12). Known from Höwenegg in Germany, Samos Main Bone Beds and White Sands (common), Mytilinii 4, Mytilinii 1A, B and C, Pikermi (rare), Ravin des Zouaves and Vathylakkos 2 and 3 in Greece. Also known from Grebeniki in the Ukraine, Cobanpinar and Kemiklitepe in Turkey and Middle and Upper Maragheh in Iran. There are several unpublished postcranial elements from Samos, indicating an animal with relatively gracile limbs (Fig. 2). Genus Lycyaena Hensel This genus, essentially Asian in distribution, is apparently the oldest member of the clade that links it with the genera Hyaenictis and Chasmaporthetes (Werdelin and Solounias, 1991), based on material from the Beglia Formation locality of Bled ed Dourah in Tunisia thought to be of early Vallesian (MN9) age (Werdelin and Turner, 1996a). The three genera are placed in Ecomorph Group 4, characterised as cursorial meat and bone eaters in which the postcarnassial molars are reduced or absent, anterior premolar cusps increase in size and the known postcranial skeleton exhibits increasingly cursorial tendencies. Lycyaena chaeretis (Gaudry) Turolian (MN11–12). Known from Pikermi and Samos Main Bone Beds in Greece and Cerro de la Garita and El Arquillo in Spain. Also known from Taraklia in the Ukraine. This species is yet another taxon with a long history of name changes and Werdelin and Solounias (1991) suggest that it may even be synonymous with the Chinese Lycyaena dubia, although it would then be the senior synonym. Genus Hyaenictis (Gaudry) This second member of Ecomorph group 4 is a poorlyknown genus that appears to have had a largely southern

distribution with its main known occurrence at Langebaanweg in South Africa. Hyaenictis graeca (Gaudry) Turolian (MN11–12). Known from Pikermi in Greece. This is the type species of the genus, although it is poorly known, owing to the lack of material. Hyaenictis almerai Villalta Comella and Crusafont Pairo´ Turolian (MN12). Known from San Miquel del Taudell in Spain. This, a possible second species of Hyaenictis, was proposed by Villalta Comella and Crusafont Pairó (1945), but Werdelin and Solounias (1991) argue that further study is required to verify this generic allocation for a specimen with a distinct suite of characters that do not sit comfortably in Hyaenictis, although they place it at that general grade of development. Genus Lycyaenops Kretzoi Werdelin (1999) has argued that this genus, named by Kretzoi (1938) but inadequately characterized by him, is valid and that it is phylogenetically close to the Lycyaena– Chasmaporthetes lineage, placed between Hyaenictis and Chasmaporthetes. Lycyaenops rhomboideae Kretzoi Ruscinian (MN14). Known from Pestlörinc in Hungary (Werdelin, 1999). Genus Chasmaporthetes Hay Skeletal features of the genus, taken in conjunction with the narrow and trenchant premolars, amply support the interpretation of a cursorial and essentially flesh-eating lifestyle and stand in marked contrast to the bone-consuming adaptations of living species of spotted, brown and striped hyaenas. Chasmaporthetes lunensis (Del Campana) Ruscinian-Villafranchian (MN14–17). Known from Etouaires, Pardines, Roccaneyra, Sénèze and St Vallier in France, Erpfinger Höhle, Neulingen and Schernfeld in Germany, Inferno, Olivola and Triversa in Italy, El Rincón, La Puebla de Valverde, Layna, Villaroya and Las Higueruelas in Spain and Dafnero in Greece. Also known from Gülyazi and Çalta in Turkey and the Odessa Catacombs in the Ukraine. The recent re-identification of the material from the German Lower Pleistocene site of Schernfeld as Chasmaporthetes lunensis rather than Pliocrocuta perrieri (Qiu, 1987) makes this the latest occurrence of the species and of the genus in Europe (Fig. 3). Recent detailed study of a well-preserved cranium from La Puebla de Valverde (Antón et al., 2007) confirms that this species was an active predator and very probably a social hunter, with a skull morphology suggesting broad functional similarity with that of the African hunting dog, Lycaon pictus.

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Chasmaporthetes borissiaki (Khomenko) Ruscinian (MN15). Known from Perpignan in France. ‘Chasmaporthetes’ bonisi (Koufos) Turolian (MN11–13). This species was originally reported from Dytiko, Greece by Koufos (1987). Werdelin and Solounias (1990, 1991) disputed this identification, referring the material in question to Adcrocuta instead. Subsequent work at this locality indeed suggests the presence of a taxon identical or closely related to Chasmaporthetes at the site (De Bonis and Koufos, 1994; Koufos, 2000). However, the issue of the taxonomic allocation of the type specimen of C. bonisi remains, since it is more like Adcrocuta than any other specimen referred to C. bonisi, calling in question not the presence of such a taxon at Dytiko but rather the species C. bonisi. However, we will accept this designation here. Synonymy with C. exitelus Kurtén and Werdelin from China (Kurtén and Werdelin, 1988) is implied, but not demonstrated. Chasmaporthetes melei Rook, Ferretti, Arca and Tuveri This is a Late Pliocene to Early Pleistocene species, endemic to Sardinia, where it has been found in the Monte Tuttavista fissure fillings (Rook et al., 2004). It is the smallest known species of Chasmaporthetes, which may be due to dwarfing in an island setting. Genus Metahyaena Viranta and Werdelin This is the most primitive genus in the Ecomorph 5 group, in which it is placed only because the premolars show in incipient form from the more bulbous shape that they have in Belbus and later Ecomorph 5 taxa. In other features, it is very similar to Ecomorph 3 taxa. Metahyaena confector Viranta and Werdelin This, the only known species of Metahyaena is from the MN9, Locality 12 of the Sinap Formation, where a single well preserved right mandibular ramus was found (Viranta and Werdelin, 2003). Genus Belbus Werdelin and Solounias Belbus, together with the genera Palinhyaena Qiu, Huang and Guo, Leecyaena Young and Liu and Ikelohyaena Werdelin and Solounias, has been placed within Ecomorph Group 5. Palinhyaena and Leecyaena are Asian in distribution, while Ikelohyaena is known only from Africa (Werdelin and Solounias, 1991; Werdelin et al., 1994; Werdelin and Turner, 1996a). However, renewed study (Werdelin pers. obs.) has shown that Leecyaena is based on a somewhat aberrant specimen of Pliocrocuta. Together, the members of these genera are referred to as transitional bone-cracking hyaenas, since they are the first within the phylogeny to exhibit adaptations towards the modern level of bone-consuming abilities.

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Belbus beaumonti (Qiu) Turolian (MN11–12). Known from Samos in Greece and also from Çobanpinar in Turkey. This is the sole known occurrence of the species, although the material has a moderately long and complex synonymy since its first description as Hyaena sp. by De Beaumont (1968). Genus Adcrocuta Kretzoi This is the first of the hyaenas to show adaptations of Ecomorph Group 6 type, the modern, fully bone-cracking morphology. The relatively complex taxonomic history of this genus has been summarised by Howell and Petter (1985) and Werdelin and Solounias (1990, 1991), who have argued that it is in fact the sister taxon of Crocuta (but see Wagner (1998) for a different view). These analyses, coupled with the referral of Adcrocuta australis at Langebaanweg to Chasmaporthetes (Werdelin and Turner, 1996a), show that only one species, A. eximia, belongs in the genus. Adcrocuta eximia (Roth and Wagner) Vallesian-Turolian (MN10–13). Widely known from various localities, including Kalimantsi and Hadgimidovo in Bulgaria, Titov Veles in FYROM, Mont Luberon in France, Dytiko, Halmyropotamos, Pikermi, Prokoma, Ravin de la Pluie, Ravin des Zouaves 1 and 5, Vathylakkos 3, Prochoma and Samos Main Bone Beds and White Sands and Xirochori 1 in Greece, Baltavar, Csákvár and Polgárdi in Hungary, Çimislia in Rumania, Dorn Durkheim in Germany, Kohfidisch in Austria and Arquillo de la Fontana, ?Concud, Los Aljezares, Los Mansuetos, Masia del Barbo, Peña del Macho and Piera in Spain. Also known from Belka, Taraklia and Grebeniki in the Ukraine, Kalmakpaj and Pavlodar in Kazakhstan, Karain, Kemiklitepe A-B, Kinik, Küçükyözgat and Upper Kavakdere in Turkey, Upper and Middle Maragheh in Iran. This species is characterised by derived dental features in the form of well-developed, conical premolars and a reduced protocone on the upper carnassial, somewhat oddly matched by the retention of P1. Postcranial remains from Pikermi show it to have been somewhat stocky in build, a powerful animal, but perhaps not particularly fast-moving. It appears suddenly at the beginning of MN10 and disappears just as suddenly at the end of MN13. Its antecedents are unknown (Fig. 2). Genus Allohyaena Kretzoi A poorly-known genus again erected by Kretzoi (1938) and again containing only one species. Allohyaena kadici Kretzoi Early Turolian (MN11). Known from Esterhézy Cave, Csákvár in Hungary and Dorn-Dürkheim in Germany. This very large species was tentatively removed from the Hyaenidae by Werdelin and Solounias (1991), but, more recently, reinstated by Werdelin and Kurtén (1999). Identification at only two sites, with the Dorn-Dürkheim material consisting

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only of two damaged premolars (Morlo, 1997), led Werdelin and Kurtén to suggest that its geographic range has not been adequately sampled. It appears to fall close in morphology to the large, bone-cracking hyaenas of Ecomorph Group 6 and Werdelin and Kurtén suggest a possible close relationship to Adcrocuta eximia. Genus Pliocrocuta Kretzoi Werdelin and Solounias (1991) discuss the taxonomic history of material now referred to this genus and, together with Turner (1990), summarise the arguments for synonymy of taxa, such as P. pyrenaica, Hyaena prisca and H. donnezani within the single species P. perrieri. As we mentioned above, it is also likely that Leecyaena is a synonym of this genus. All of the material, whatever its taxonomic affinities, clearly falls within Ecomorph Group 6. Pliocrocuta perrieri Croizet and Jobert Ruscinian-Post Villafranchian (MN15–18 and later). Known widely from localities, such as Hollabrunn in Austria, Ardé, Etouaires, Es-Taliens, Chilhac, L’Escale, Lunel Viel, Montmaurin, Montsaunes, Pardines, Perpignan, Sénèze, Serrat-d’En-Vacquer, St-Vallier and Vialette in France, Erpfinger Höhle, Greusnach, Gundersheim 1 and 4, Mauer and Mosbach in Germany, Hajnácka in Slovakia, Red Crag deposits in England, Gerakarou, Sesklon and Petralona in Greece, Montopoli and Tasso in Italy, Tegelen in the Netherlands, Furninha in Portugal and La Calera II, El Rincón, La Gloria 4, La Puebla de Valverde, Layna and Villarroya in Spain. This species has a very disjointed pattern of appearance in Europe. It first seems to have become locally extinct towards the end of the Pliocene, after its appearance at Puebla de Valverde, Chilhac, Tegelen and a number of localities in the Red Crag deposits of Suffolk (Turner, 1995a). It reappeared in the earlier stages of the Middle Pleistocene at sites, such as Petralona, Mauer and Mosbach 2 and it is also known from the southern French sites of L’Escale and Lunel Viel where it has previously been identified as a large striped hyaena, Hyaena prisca De Serres (Turner, 1990; Werdelin and Solounias, 1991) (Fig. 3). This second dispersion of P. perrieri seems to have been somewhat restricted, since it does not appear to have reached Italy or Britain (Turner, 1995a, 1995b). However, it may have reached the Iberian Peninsula, contrary to earlier suggestions (Turner, 1995b), in view of the presence there of what has been identified as Hyaena prisca at the Portuguese Upper Pleistocene locality of Furninha (Cardoso, 1992). We have not seen the material, but it appears to us, based on Cardoso’s descriptions, measurements and illustrations, to be identical to other specimens of what we regard as P. perrieri. Whether this material actually represents a population of P. perrieri and, if so, whether it implies a recolonisation or a relict Villafranchian population are both unclear to us at present.

Genus Pachycrocuta Kretzoi Kretzoi erected the genus Pachycrocuta in 1938 to accommodate the gigantic short-faced hyaena, P. brevirostris. Previous discussions by many authors have placed Pliocrocuta perrieri within this genus, but the cladistic analyses of Werdelin and Solounias (1991) show them as only sister taxa and that in only five out of 16 trees. Pachycrocuta brevirostris (Aymard) Villafranchian to Middle Pleistocene. Known from Stranska Skala in the Czech Republic, Bacton, Mundesley, Sidestrand and Westbury-sub-Mendip in England, Sainzelles, Ceyssaguet and Vallonnet in France, Süssenborn, Untermassfeld and Wurzburg-Schalksberg in Germany, Gerakarou, Appollonia and Petralona in Greece, Gombaszoeg in Hungary, Olivola, Casa Frata and Pirro Nord in Italy, Cueva Victoria, Venta Micena, Almenara 1, Ponton de la Oliva, Barranco León 5, Fuenta Nueva 3 and Incarcal in Spain. This hyaena was first recorded from Sainzelles in the Auvergne (Aymard, 1846) and described in detail by Boule (1893). The species, the largest true hyaena known and a member of Ecomorph Group 6 par excellence (Fig. 3), is widely distributed in Eurasian Lower and Middle Pleistocene localities. It was discussed in some detail by Howell and Petter (1980) and has been comprehensively reviewed by Turner and Antón (1996) who discussed its likely bone-accumulating abilities (Fig. 3). The animal is best represented at Zoukoudian in China, where Pei (1934) estimated the remains of some two thousand individuals were represented. Elsewhere, and particularly in Europe, it is usually represented by a small number of specimens and individuals, the most notable exception being the German later Lower Pleistocene locality of Untermassfeld, where at least 18 animals are currently recorded (Turner, 2001). Genus Crocuta Kaup The spotted hyaena, Crocuta crocuta (Erxleben) is today the sole species referred to the genus Crocuta. Now confined to Africa (Kruuk, 1972; Mills, 1990), where it is widely distributed in the south of the Sahara. It was formerly present in much of Eurasia during the Pleistocene, appearing in what can only be described as superabundance in many western European Upper Pleistocene cave deposits (Kurtén, 1968) to the point where a listing of sites would be enormous. Its earliest-known appearance in Europe is at Atapuerca, Spain, dated c. 0.78 Ma (Garcia and Arsuaga, 2001). It was particularly common in British deposits of last glacial and last interglacial age (Turner, 1995a). As Werdelin and Solounias (1991) pointed out, opinions over the question of taxonomic separation of the fossil material have varied greatly, with some authors referring the European form to the species Crocuta spelaea, although the known plasticity of the living species has led other authors to argue for similar effects in the past (Kurtén, 1957; Turner, 1984, 1990). The European form, whatever its precise taxonomic status, appears to have been a prodigious collector of bones in many areas (Fig. 3).

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3. Evolutionary patterns among the European Hyaenidae The earliest record of the Hyaenidae is in MN4, as pointed out by Werdelin and Solounias (1996), although the patchy nature of the earliest records makes it difficult to be certain about the continent of origin (Werdelin and Turner, 1996a). What is much clearer, despite the shortcomings of the fossil record, is the overall pattern of morphological evolution in the family over that long timespan, from the earliest viverrid- and herpestid-like forms through dog-like and more cursorial taxa to the larger, bone-crunching animals of the later Miocene and the Pliocene–Pleistocene epochs (Fig. 2 and 3). As we have previously pointed out (Werdelin and Turner, 1996b), it has long been recognised that there is a considerable turnover in the terrestrial mammalian fauna of Europe at the Miocene–Pliocene boundary. Savage and Russell (1983) listed some 122 of the 178 known later-Miocene genera failing to appear in the Pliocene, with the Carnivora reduced from 59 later-Miocene species to only 32 in the Early Pliocene. The Hyaenidae stand out within this larger pattern because of the evolution in Ecomorph Groups. This pattern, paralleled across the whole of the Old World (Werdelin and Turner, 1996a, 1996b; Turner and Antón, 2004), is particularly evident at the end of the Miocene and during the Pliocene, where the reduction in canid-like morphotypes among the hyaenas is eventually matched by the rise in numbers and ecological importance of the Canidae. During the latest Miocene, almost, the only hyaenid to exhibit the bone-smashing capabilities of the living Ecomorph Group 6 species is Adcrocuta eximia. The poorly-known Allohyaena kadici would, of course, also fall within the same group as Adcrocuta eximia, but it is so far known only from one Hungarian and one German site and it is difficult to attach greater significance to this material at the moment. However, if two taxa are recognised, this lends more credence to the idea that latest Miocene hyaenas were undergoing a major transition in their way of life. Adcrocuta eximia has been characterised as powerful, but perhaps rather slow-moving, with implications that it was more of a specialist scavenger (Werdelin and Turner, 1996a, 1996b), but this view may warrant revision. In the first place, its upper carnassial is rather sectorial, with a reduced protocone and may imply a rather high-flesh content in the diet. Secondly, its great size not only indicates its ability to appropriate carcasses, but also implies a clear advantage when seeking to capture medium to large-sized ungulates, as discussed by Turner and Antón (1996). Among modern Hyaenidae, the extent of aggressive behaviour appears to be conditioned by dominant relationships between large carnivores in general and between sympatric hyaenas in particular. Striped hyaenas living in areas where they are not subject to competition from spotted hyaenas are larger and tend to prey more actively on large ungulates (Rieger, 1981; Kingdon, 1990). Kowalski and RzebikKowalska (1991) report a single individual striped hyaena in Algeria, killing and eating 12 asses in one season. As by far the largest hyaena in its habitat Adcrocuta eximia was probably able not only to expropriate carcasses from other predators but

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also to defend its own kills, although group action would undoubtedly have been necessary in any disputes involving the enormous Machairodus giganteus. A third point concerns the subject of cursoriality directly. In earlier descriptions of Adcrocuta eximia as a ‘‘lumbering’’ scavenger (Werdelin and Solounias, 1996) it is implied that the animal could not run fast. However, the postcrania from Pikermi, described by Gaudry (1861) and examined by us in Paris, appear no more robust than those of European Crocuta crocuta or of Pachycrocuta brevirostris and both of those are unlikely to have been exclusively scavengers (Turner and Antón, 1996). In any efforts to characterise fossil species we must beware of tendencies to pigeon-holing of taxa and it is clear that scavenging and hunting are simply part of a spectrum of behaviours and that living hyaenas show a range of foodobtaining strategies that crosscut various aspects of their skeletal morphology. A relevant point here is the likely ecology and behaviour patterns of the Miocene dog-like hyaenas, which clearly warrants further investigation. Although Martin (1989) has suggested that social hunting by packs is an entirely PlioPleistocene development, linked to the presence of large ungulate herds in open grasslands, we suspect that this view may be too simplistic. Our image of predator–prey relationships is strongly influenced by studies of eastern African carnivores, easily observed in high-visibility parks, such as the Mara and Serengeti. Such areas are probably not the best analogues for the seasonal woodlands of the Turolian of Western Europe. A more appropriate comparison can be made with the hunting behaviour of the dhole (Cuon alpinus), which operates as a pack hunter of medium to large-sized prey in the woodlands and forests of present-day India. In spite of being considerably smaller than an average wolf and of having a skeleton only moderately adapted for cursorial locomotion, the dhole is a very efficient group-living pursuit hunter and killer. The Pliocene sees the full emergence of a differently organised guild of large carnivores in Europe, with machairodont cats, largely specialised as flesh-eaters, dependent on seizing prey and holding it still while killing it in order to avoid (or at least minimise) damage to their canines (Werdelin and Turner, 1996b; Turner and Antón, 1997; Salesa et al., 2005, 2006). The hyaenas have by then started to show the specialised ability to demolish bone with the appearance of Pliocrocuta perrieri in place of Adcrocuta eximia and thus, were able to obtain a consistent living from scavenging when necessary, although the Ecomorph Group 4-hunting hyaenas of the genus Chasmaporthetes continue to operate successfully until the end of the Pliocene. By the earliest Pleistocene, however, the only hyaena still operating in Europe, with the possible, implied exception of a remnant population of Pliocrucota perrieri in Portugal, was the gigantic Pachycrocuta brevirostris (Turner, 1992, 1995a, 1995b, 1995c). Whether the reappearance of Pliocrucota perrieri in Europe during the Middle Pleistocene represents a recolonisation from Asia or a dispersion out from Portugal has been commented on by us before (Turner and Antón, 1996), since it is unclear whether the Upper Pleistocene occurrence of the species at Furninha Cave is a late remnant of

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this second European appearance or a longer standing relict of the original Villafranchian population. However, its absence from elsewhere in the Iberian Peninsula after its record at earlier Villafranchian localities in Spain tends to argue against movement out from Portugal during the Middle Pleistocene. In the Middle Pleistocene, we see the final turnover before the appearance of fully modern fauna of the Holocene when hyaenas finally disappeared from Europe. Pachycrocuta brevirostris and Pliocrocuta perrieri, together with the sabre-toothed cat (Homotherium latidens), the cheetah (Acinonyx pardinensis) and the European ‘‘jaguar’’ (Panthera gombaszoegensis), became extinct. In their place, Europe was left with a large predator guild consisting of Crocuta crocuta, which had arrived by the latter part of the Lower Pleistocene to be followed (or perhaps accompanied, the data are not really clear) by lion, Panthera leo and leopard, Panthera pardus (Turner, 1995c; Garcia and Arsuaga, 1999; Antón et al., 2005). In addition, as pointed out by Turner and Antón (1996), the various species of Canis that appeared in increasing numbers during the Pleistocene probably operated as pack-hunting animals able to harry other, larger predators and seize carcasses. Canis etruscus and/or Canis mosbachensis remains are found in considerable numbers at many Lower and Middle Pleistocene sites, perhaps indicating large local populations. Canis (Xenocyon) falconeri and Canis (X.) lycaonoides, species approaching the size of a modern wolf, may also have added to the pressure on other taxa in direct competition for carcasses and undertaken their own share of body-part accumulation. Thus, while Crocuta crocuta was to become the local bone accumulator par excellence, it is unlikely to have been the first. On grounds of size and morphology, it would seem entirely likely that Pliocrocuta perrieri and Pachycrocuta brevirostris, and very probably Adcrocuta eximia, Allohyaena kadici and perhaps Belbus beaumonti, were among the hyaenas that shared some of their abilities with the various canids. Any European Plio-Pleistocene locality may therefore, at least potentially, contain a hyaena-accumulated bone assemblage, a trap awaiting the unwary archaeologist seeking evidence of earliest human presence in the patterning of such assemblages in the absence of unequivocal artefacts or human skeletal remains. Acknowledgement We thank numerous people in many institutions for access to specimens in their care and for their ready assistance. References Alcala, L., 1994. Macromamíferos Neógenos de la Fosa de Alfambra-Teruel 3. Museo Nacional de Ciencias Naturales, Madrid. Alcala, L., Morales, J., Soria, D., 1992. El registro fósil neógeno de los carnívoros (Creodonta y Carnivora, Mammalia) de España. Paleontología i Evolució 23, 55–66. Antón, M., Galobart, A., Turner, A., 2005. Co-existence of scimitar-toothed cats, lions and hominins in the European Pleistocene. Implications of the post-cranial anatomy of Homotherium latidens (Owen) for comparative palaeoecology. Quaternary Science Reviews 24, 1287–1301.

Antón, M., Turner, A., Salesa, M.J., Morales, J., 2007. A complete skull of Chasmaporthetes lunensis (Carnivora, Hyaenidae) from the Spanish Pliocene site of La Puebla de Valverde (Teruel). Estudios Geológicos 62, 375–388. Aymard, A., 1846. Communication sur le gisement de Sainzelles. Annales de la société d’agriculture, science et arts du Puy 12, 153–155. Boule, M., 1893. Description de l’Hyaena brevirostris du Pliocène de Sainzelles près de Le Puy (Haute-Loire). Annales des sciences naturelles 15 (série 8), 85–97. Cardoso, J.L., 1992. Contribuição Para o Conhecimento dos Grandes Mamiferos do Pleistocénico Superio de Portugal. Thesis, Universidade Nova de Lisboa, Portugal (unpublished). Crusafont Pairó, M., Petter, G., 1969. Contribution à l’étude des Hyaenidae. La sous-famille des Ictitheriinae. Annales de paléontologie (vertèbres) 55, 89–127. De Beaumont, G., 1968. Une intéressante mandibule de Hyaenidae (Carnivora) du Pontien de Samos. Archives des sciences de Genève 22, 21–26. De Beaumont, G., 1986. Les carnivores (mammifères) du Néogène de Höwenegg/Hegau, Baden-Württemberg. Carolinea 44, 35–45. De Bonis, L., 1994. Carnivores viverroïdes (Carnivora, Mammalia) du Miocène inférieur d’Europe occidentale : origine des familles modernes de Feliformia. Bulletin de la Société géologique de France 165, 85–92. De Bonis, L., Koufos, G., 1994. Some Hyaenidae from the Late Miocene of Macedonia (Greece) and a contribution to the phylogeny of the hunting hyaenas. Münchner Geowissenschaftliche Abhandlungen 26, 81–96. Garcia, N., Arsuaga, J.L., 1999. Carnivores from the Early Pleistocene hominidbearing Trinchera Dolina 6 (Sierra de Atapuerca, Spain). Journal of Human Evolution 37, 415–430. Garcia, N., Arsuaga, J.L., 2001. Les carnivores (Mammalia) des sites du Pléistocène ancien et moyen d’Atapuerca (Espagne). L’Anthropologie 105, 83–93. Gaudry, A., 1861. Note sur les carnassiers fossiles de Pikermi (Grèce). Bulletin de la Société géologique du France 18 (série 2), 527–537. Gaudry, A., 1862–1867. Animaux fossiles et géologie de l’Attique. Éditions F. Savy, Paris. Howell, F.C., Petter, G., 1980. The Pachycrocuta and Hyaena lineages (PlioPleistocene and extant species of the Hyaenidae). Their relationships with Miocene Ictitheres: Palhyaena and Hyaenictitherium. Geobios 13, 579– 623. Howell, F.C., Petter, G., 1985. Comparative observations on some Middle and Upper Miocene hyaenid genera: Percrocuta Kretzoi, Allohyaena Kretzoi, Adcrocuta Kretzoi (Mammalia, Carnivora, Hyaenidae). Geobios 18, 419–476. Kowalski, K., Rzebik-Kowalska, B., 1991. Mammals of Algeria. Polish Academy of Sciences, Warsaw. Kingdon, J., 1990. Arabian mammals: a natural history. Academic Press, London. Koepfli, K.-P., Jenks, S.M., Eizirik, E., Zahirpour, T., van Valkenburgh, B., Wayne, R.K., 2006. Molecular systematics of the Hyaenidae: relationships of a relictual lineage resolved by a molecular supermatrix. Molecular Phylogenetics and Evolution 38, 603–620. Koufos, G.D., 1987. Chasmaporthetes bonisi, a new hyaenid (Carnivora, Mammalia) from the late Miocene of Macedonia (Greece). Bulletin de la Société géologique du France 3 (8), 913–920. Koufos, G., 2000. Revision of the late Miocene carnivores from the Axios valley, Macedonia, Greece. Münchner Geowissenschaftliche Abhandlungen 39, 51–92. Kretzoi, M., 1938. Die Raubtiere von Gombaszög, nebst einen Übersicht der Gesamtfauna. Annales Museum Nationale Hungaricum 31, 88–137. Kretzoi, M., 1951. A Csákvári Hipparion-fauna. Földtani Közlön 81, 384–417. Kretzoi, M., 1952. Die Raubtiere der Hipparionfauna von Polgárdi. Annales Instituti Geologici Hungarici 40, 1–42. Kretzoi, M., 1984. Felsö pannóniai. A sümeg–gerinci fauna és faunaszakasz. Geologica Hungarica, serie geologica 20, 214–222. Kruuk, H., 1972. The spotted hyaena. University of Chicago Press, Chicago. Kurtén, B., 1957. The bears and hyaenas of the interglacials. Quaternaria 4, 69–81. Kurtén, B., 1968. Pleistocene mammals of Europe. Weidenfeld and Nicolson, London.

A. Turner et al. / Geobios 41 (2008) 677–687 Kurtén, B., Werdelin, L., 1988. A review of the genus Chasmaporthetes Hay, 1921 (Carnivora, Hyaenidae). Journal of Vertebrate Paleontology 8, 46–66. Martin, L.D., 1989. Fossil history of the terrestrial carnivora. In: Gittleman, J.L. (Ed.), Carnivore behaviour, ecology and evolution. Chapman and Hall, London, pp. 536–568. Mills, G., 1990. Kalahari hyaenas. Unwin Hyman, London. Morlo, M., 1997. Die Raubtiere (Mammalia, Carnivora) aus dem Turolium von Dorn-Dürkheim 1 (Rheinhessen, Deutschland). Teil 1: Mustelidae, Hyaenidae, Percrocutidae, Felidae. Courier Forschungsinstitut Senckenberg 197, 11–47. Pei, W.C., 1934. On the carnivora from Locality 1 of Choukoutien. Palaeontologica Sinica ser C 8 (1), 1–217. Pilgrim, G., 1931. Catalogue of the Pontian Carnivora of Europe in the Department of Geology. British Museum (Natural History), London. Qiu, Z., 1987. Die Hyaeniden aus dem Ruscinium und Villafranchium Chinas. Münchner Geowissenschaftlichen Abhandlungen A 9, 1–108. Rieger, I., 1981. Hyaena hyaena. Mammalian Species 150, 1–5. Rook, L., Ferretti, M.P., Arca, M., Tuveri, C., 2004. Chasmaporthetes melei n. sp., an endemic hyaenid (Carnivora, Mammalia) from the Monte Tuttavista fissure fillings (Late Pliocene to Early Pleistocene; Sardinia, Italy). Rivista Italiana di Paleontologia e Stratigrafia 110, 707–714. Salesa, M.J., Antón, M., Turner, A., Morales, J., 2005. Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bite. Zoological Journal of the Linnean Society 144, 363–377. Salesa, M.J., Antón, M., Turner, A., Morales, J., 2006. Inferred behaviour and ecology of the primitive sabre-toothed cat Paramachairodus ogygia (Felidae, Machairodontinae) from the Late Miocene of Spain. Journal of Zoology 268, 243–254. Savage, D.E., Russell, D.E., 1983. Mammalian paleofaunas of the world. Addison-Wesley Publishing Co, London. Semenov, Y.A., 1985. Ictitherium pannonicum (Carnivora, Viverridae) from Meiotic deposits of the northern Black Sea area. Vestnik Zoologii 23–27 (in Russian). Semenov, Y.A., 1989. Ictitheres and morphologically related hyaenas from the Neogene of the USSR. Naukova Dumka, Kiev (in Russian). Thenius, E., 1966. Zur Stammesgeschichte der Hyänen (Carnivora, Mammalia). Zeitschrift für Säugetierkunde 31, 293–300. Turner, A., 1984. The interpretation of variation in fossil specimens of spotted hyaena (Crocuta crocuta Erxleben, 1777) from Sterkfontein Valley sites (Mammalia: Carnivora). Annals of the Transvaal Museum 33, 399–418. Turner, A., 1990. The evolution of the guild of larger terrestrial carnivores in the Plio-Pleistocene of Africa. Geobios 23, 349–368. Turner, A., 1992. Villafranchian-Galerian larger carnivores of Europe; dispersions and extinctions. In: von Koenigswald, W., Werdelin, L. (Eds.), Mammalian migration and dispersal events in the european quaternary. Courier Forschungsinstitut Senckenberg 153, pp. 153–160. Turner, A., 1995a. Evidence for Pleistocene contact between the British Isles and the European continent based on distributions of larger carnivores. Geological Society Special Publication 96, 141–149. Turner, A., 1995b. Regional variations in Lower and Middle Pleistocene larger mammal faunas of Europe: an Iberian perspective. In: Bermúdez de Castro,

687

J.M., Carbonell, E., Arsuaga, J.L. (Eds.), Evolución humana en Europa y los yacimientos de la Sierra de Atapuerca. Junta de Castilla y León, Valladolid, pp. 57–73. Turner, A., 1995c. The Villafranchian large carnivore guild: geographic distribution and structural evolution. Il Quaternario 8, 349–356. Turner, A., 2001. Remains of Pachycrocuta brevirostris (Mammalia, Hyaenidae) from the Lower Pleistocene site of Untermassfeld near Meiningen. In: Kahlke, H.D., Kahlke, R.D. (Eds.), Das Pleistozän von Untermassfeld in Thüringen. Rudolf Harbelt Verlag, Bonn, pp. 673–690. Turner, A., Antón, M., 1996. The giant hyaena, Pachycrocuta brevirostris (mammalian, carnivora, Hyaenidae). Geobios 29, 455–468. Turner, A., Antón, M., 1997. The big cats and their fossil relatives: an illustrated guide to their evolution and natural history. Columbia University Press, New York. Turner, A., Antón, M., 2004. Evolving eden: an illustrated guide to the evolution of the African large mammal fauna. Columbia University Press, New York. Villalta Comella, J.F., Crusafont Pairó, M., 1945. Nuevas aportaciones al conocimiento de los Carnívoros Pontienses del Vallé Penedés. Publicaciones del Instituto Geológico, Diputacion Provincial de Barcelona 7, 81–121. Viranta, S., Werdelin, L., 2003. Carnivora. In: Fortelius, M., Kappelman, J., Sen, S., Bernor, R.L. (Eds.), Geology and Paleontology of the Miocene Sinap Formation, Turkey. Columbia University Press, New York, pp. 178– 193. Wagner, P.J., 1998. A likelihood approach for evaluating estimates of phylogenetic relationships among fossil taxa. Paleobiology 24, 430–449. Werdelin, L., 1999. Studies of fossil hyaenids: the identity of Lycyaenops rhomboideae Kretzoi from Pesztlörinc, Hungary. Zoological Journal of the Linnean Society 126, 307–317. Werdelin, L., Kurtén, B., 1999. Allohyaena (Mammalia: carnivora): giant hyaenid from the Late Miocene of Hungary. Zoological Journal of the Linnean Society 126, 319–334. Werdelin, L., Solounias, N., 1990. Studies of fossil hyaenids: the genus Adcrocuta Kretzoi and the interrelationships of some hyaenid taxa. Zoological Journal of the Linnean Society 98, 363–386. Werdelin, L., Solounias, N., 1991. The Hyaenidae: taxonomy, systematics and evolution. Fossils and Strata 30, 1–104. Werdelin, L., Solounias, N., 1996. The evolutionary history of hyenas in Europe and western Asia during the Miocene. In: Bernor, R.L., Fahlbusch, V., Rietschel, S. (Eds.), Later Neogene European biotic evolution and stratigraphic correlation. Columbia University Press, New York, pp. 290–306. Werdelin, L., Turner, A., 1996a. The fossil Hyaenidae of Africa: present status. In: Stewart, K.M., Seymour, K.L. (Eds.), Palaeoecology and palaeoenvironments of Late Cenozoic mammals. University of Toronto Press, Toronto, pp. 637–659. Werdelin, L., Turner, A., 1996b. Turnover in the guild of larger carnivores in Eurasia across the Miocene–Pliocene boundary. Acta Zoologica Cracoviensia 39, 585–592. Werdelin, L., Turner, A., Solounias, N., 1994. Studies of fossil hyaenids: the genera Hyaenictis Gaudry and Chasmaporthetes Hay, with a reconsideration of the Hyaenidae of Langebaanweg, South Africa. Zoological Journal of the Linnean Society 111, 197–217.