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Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe

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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.

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... In many parts of the world including Europe, carnivore assemblages were strongly impacted by Late Quaternary extinction trajectories (Stuart and Lister 2007;Malhi et al. 2016;Galetti et al. 2018), putting a final nail to the coffin of most large-bodied carnivores and socalled mega-carnivores that are a hallmark of Pleistocene animal ecologies (Faith et al. 2019). The Late Pleistocene in particular has seen substantial turnover in carnivore community dynamics and composition and documents the extinction of some notable carnivore others (Turner 1985), such as various hyena species with whom hominins in Europe and elsewhere have co-evolved and shared the environment for many millennia (Turner et al. 2008). Some scholars have argued that Eurasian Neanderthals in particular are to be understood as an integral part of situated carnivore ecologies and their associated predator-prey interlinkages, whose gradual disappearance in the second part of the Late Pleistocene may have consequently precipitated the eventual demise of the Neanderthal phenotype at the end of MIS 3 (Estévez 2004;Stewart 2005Stewart , 2007. ...
... This is astonishing giving the well-documented, close ecogeographic association of early hominins and various hyena taxa throughout the Lower and Middle Pleistocene, especially in Africa (Turner 1990;Werdelin and Lewis 2005;Treves and Palmquist 2007). Even though the involved taxonomic groups change over time (Turner et al. 2008), Hominidae and Hyaenidae share an exceptionally long evolutionary history (Brantingham 1998;Stiner 2012), and either of the two is often a strong proxy for the presence of the respective other, as reflected in early expansions of faunal assemblages with hominins and hyenas from Africa into the Eastern Mediterranean and Western Europe (Arribas and Palmqvist 1999;Lewis and Werdelin 2010;Brugal et al. 2020), for example documented at key sites such as Dmanisi (Coil 2016), Pakefield (Parfitt et al. 2005;Iannucci et al. 2021) and Atapuerca (García and Arsuaga 2001). ...
... The ancestral cousins of today's spotted hyenas (Crocuta crocuta) arrived in Europe shortly before 300 kya Turner et al. 2008), broadly falling into the second half of the Middle Pleistocene and thus predating 'classic' fossil Neanderthal phenotypes which only appear around MIS 5 (Serangeli and Bolus 2008). This timing bears evolutionary significance because what we recognize as Neanderthal biocultural characteristics developed gradually between ca. ...
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Late Pleistocene hominins co-evolved with non-analogue assemblages of carnivores and carnivorous omnivores. Although previous work has carefully examined the ecological and adaptive significance of living in such carnivore-saturated environments, surprisingly little attention has been paid to the social and cultural consequences of being-with, and adapting to, other charismatic predators and keystone carnivores. Focusing on Neanderthal populations in Western Eurasia, this paper draws together mounting archaeological evidence that suggests that some Late Pleistocene hominins devised specific behavioral strategies to negotiate their place within the vibrant carnivore guilds of their time. We build on integrative multispecies theory and broader re-conceptualizations of human-nature relations to argue that otherwise puzzling evidence for purported ‘symbolic’ behavior among Neanderthals can compellingly be re-synthesized with their ecology, settlement organization and lifeworld phenomenology. This re-framing of Neanderthal lifeways in the larger context of startling carnivore environments reveals that these hominins likely developed intimate, culturally mediated, and hence varied, bonds with raptor, hyena and bear others, rather than merely competing with them for resources, space and survival. This redressing of human-carnivore relations in the Middle Paleolithic yields important challenges for current narratives on evolving multispecies systems in the Late Pleistocene, complicating our understanding of Late Quaternary megafaunal extinctions and the roles of hominins in these processes.
... The family Hyaenidae Gray (1821) is represented today only by four species: Crocuta crocuta (Erxleben 1777), Hyaena hyaena (Linnaeus 1758), Parahyaena brunnea (Thunberg 1820) and Proteles cristatus (Sparrman 1783). However, the fossil record of the family includes a very wide range of body sizes and dietary adaptations (Werdelin and Solounias 1991;Turner et al. 2008;Coca-Ortega and Pérez-Claros 2019). The most primitive hyenas are small-sized, viverrid/herpestid-like genera, like Protictitherium Kretzoi (1938), Plioviverrops Kretzoi (1938) and Tungurictis Colbert (1939). ...
... Several different schemes have been proposed during the past two centuries. The scheme of Werdelin and Solounias (1991), slightly modified by Turner et al. (2008), suggests a relatively linear phylogenetic tree for the fossil Hyaenidae, with different genera gradually diverging at different ages. On the other Semenov (1989Semenov ( , 2008 identified a split of two lineages: the Ictitheriinae sensu stricto (genera Ictitherium Wagner 1848, and Thalassictis; Gervais 1850, ex Von Nordmann) and the tribe Hyaenotheriini Semenov (1989) (genera Hyaenotherium Semenov 1989Kretzoi 1938, and Miohyaenotherium;Semenov 1989). ...
... Another genus that had been considered as similar to Thalassictis is Progenetta Depéret (1892). This genus was used to include the small-sized Progenetta gaillardi Forsyth Major (1903), and Progenetta crassa (Depéret 1892) (Crusafont Pairó andPetter 1969), which are now attributed to Protictitherium (Werdelin and Solounias 1991;Turner et al. 2008;Koufos 2011;Mayda et al. 2015). Additionally it included the medium-sized Progenetta certa Forsyth Major (1903), Progenetta proava (Pilgrim 1910) and Progenetta montadai Villalta Comella and Crusafont Pairó (1943) (Viret 1951;Crusafont Pairó and Petter 1969;Crusafont Pairó and Golpe Posse 1973) that have now been attributed to Thalassictis (Werdelin and Solounias 1991;Turner et al. 2008;Mayda et al. 2015). ...
Article
The present paper deals with new hyaenid material from the locality of Hammerschmiede (Bavaria, Germany). The described specimens are attributed to two forms: most of the specimens belong to the species Thalassictis montadai, whereas one I3 is attributed to a large bone-cracking hyena. The material comes from the layers HAM 5 (11.62 Ma) and HAM 6 (slightly younger than 11.44 Ma) of Hammerschmiede (base of Late Miocene). The species Thalassictis montadai is well-known from late Aragonian and early Vallesian localities of central and southern Europe and west Asia. The presented material enables us to make a short review of the state-of-the-art about the fossil record of this species and to discuss its intraspecific variability. A gradual replacement of Thalassictis montadai, Thalassictis robusta and Hyaenictitherium wongii in Europe is demonstrated, until the arrival of canids during the latest Miocene. Additionally, the upper incisor of the large hyaenid creates some interesting questions concerning the first appearance of the crocutoid hyenas in the fossil record and their dominance over the percrocutoids.
... Hyenas are infamously known for their scavenging and carcassdismembering attitudes, with all living species, apart from the myrmechophagous Proteles cristatus (Sparrmann, 1783), regarded as "bone-crackers" (Werdelin andSolounias, 1991, 1996). This ecomorphotype become dominant after the Miocene-Pliocene transition, a period that marks a significant drop in the diversity of the family (Werdelin and Turner, 1996;Turner et al., 2008), and a general reorganization of mammalian paleocommunities (Fortelius et al., 2006;Kaya et al., 2018;Iannucci et al., 2021). ...
... Ma (Sardella and Petrucci, 2012;Madurell-Malapeira et al., 2017), P. brevirostris and C. crocuta are often considered to coexist in early Middle Pleistocene faunal assemblages of central Europe and England, with the giant hyena apparently surviving longer in these regions, up to~0.5 Ma (Kurt en, 1968;Turner and Ant on, 1996). Moreover, in this scenario another character is often overshadowed, the other bone-cracking hyena Hyaena prisca de Serres et al., 1828, whose relationship with the living striped hyena, Hyaena hyaena (Linnaeus, 1758) and brown hyena, Parahyaena brunnea (Thunberg, 1820), and the extinct Pliocene Pliocrocuta perrieri (Croizet and Jobert, 1828) are debated (e.g., Howell and Petter, 1980;Turner, 1990;Werdelin and Solounias, 1991;Turner et al., 2008). ...
... Tiraspol (Moldova) was listed among the sites with P. brevirostris by Turner and Ant on (1996), but not by Turner et al. (2008). Indeed, according to Nikiforova et al. (1971) the only Hyaenidae specimen from the site is a metopodial assigned to Crocuta sp. ...
Article
The giant, short-faced hyena Pachycrocuta brevirostris was the largest Hyaenidae ever existed and the one that perfectly embodied the distinctive bone-cracking adaptations of this mammal family. Its dispersal into Europe is regarded as a biochronological marker of the Late Villafranchian at ~2.0 Ma, and its potential ecological interactions with other carnivorans and early Homo populations diffusing Out of Africa have given rise to extensive discussions. Nevertheless, our comprehension of the extinction of P. brevirostris remains vague. Here, we first critically evaluate the European fossil record of the species and then we review the whole Epivillafranchian and Galerian Hyaenidae record, including P. brevirostris, Crocuta crocuta and "Hyaena" prisca. Biometric comparisons with other extinct and extant bone-cracking hyenas are carried out. In contrast to a common view, we recognize that there is neither evidence of a persistence of P. brevirostris beyond the Early-Middle Pleistocene boundary, nor of a coexistence between the giant hyena and C. crocuta. The replacement between the two species, which was also accompanied by the arrival of "H." prisca, occurred at ~0.8 Ma and can serve as a marker of the Epivillafranchian–Galerian turnover, part of the faunal renewal that reflects the response of mammal communities to the Early–Middle Pleistocene Transition. Moreover, we clarified that Pliocrocuta perrieri and "H." prisca were different species, and that the latter was relatively more widespread than often assumed, being recorded from localities spanning in age almost the whole Middle Pleistocene and even the early Late Pleistocene.
... However, thanks to mainly complete tooth row and moderately worn teeth, it is possible to provide a clear taxonomic affiliation. (Qiu 1987;Turner et al. 2008;A.M. pers. obs.). ...
... During the Early Pleistocene, the species reached Europe (most probably from Asia) (Werdelin 1999;Palmqvist et al. 2011). In a short time, P. brevirostris replaced two other large hyaenid species, the large bone-crushing Pliocrocuta perrieri and the gracile pack hunter Chasmaporthetes lunensis (Turner and Antón 1996;Turner et al. 2008). It was present in Europe between 2.0 and 0.7 Ma, and finally disappeared during the early Middle Pleistocene (Fig. 2; Table 2; Turner and Antón 1996;Arribas and Palmqvist 1999;Turner et al. 2008;Palmqvist et al. 2011). ...
... In a short time, P. brevirostris replaced two other large hyaenid species, the large bone-crushing Pliocrocuta perrieri and the gracile pack hunter Chasmaporthetes lunensis (Turner and Antón 1996;Turner et al. 2008). It was present in Europe between 2.0 and 0.7 Ma, and finally disappeared during the early Middle Pleistocene (Fig. 2; Table 2; Turner and Antón 1996;Arribas and Palmqvist 1999;Turner et al. 2008;Palmqvist et al. 2011). ...
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Cranial material of Pachycrocuta brevirostris from the late Early Pleistocene site of Nogaisk is the first record of this species in Ukraine. This large hyena was a representative of the Tamanian faunal complex and a single specialised scavenger in these faunas. The revisited European records list of P . brevirostris documented the presence of this species in 101 sites, dated in the range of 3.5–0.4 Ma. This species first disappeared in Africa, survived in Europe until ca. 0.8–0.7 Ma, and its last, relict occurrence was known from south-eastern Asia. The main reason of extinction of P . brevirostris probably was the competition with Crocuta crocuta . The cave hyena was smaller, but its teeth were proportionally larger to the body size, better adapted to crushing bones and slicing meat, and could also hunt united in larger groups.
... The giant short-faced hyena, Pachycrocuta brevirostris (Gervais, 1850), type species of the genus Pachycrocuta Kretzoi, 1938, was the largest and most spectacular bone-cracking hyena that ever existed (Turner and Antón, 1996;Werdelin, 1999;Alba et al., 2015). Its iconic suite of craniodental features including powerfully built skull with an inflated frontal region, wide zygoma, well-developed sagittal crest and frontal sinuses, massive premolars with microstructurally reinforced enamel , are widely considered to be adaptations for a specialist lifestyle, scavenging the kills of other sympatric large carnivores (Howell and Petter, 1980;Turner and Antón, 1996;Liu, 2001;Ferretti, 2007;Turner et al., 2008;Palmqvist et al., 2011). Similar to the extant spotted hyena (Crocuta crocuta), the short-faced hyena was able to efficiently smash large bones to access the nutritious marrow and spongy bone tissues (Kurtén, 1968;Palmqvist et al., 2011), and consequently classified within the fully-evolved bone-cracking hyaenid ecomorphotype , i.e. "morphotype 6" as defined by Werdelin and Solounias (1996). ...
... For more than a century, its authorship was generally attributed to Auguste Aymard with uncertainty about the date of publication (e.g. Kurtén, 1956;Howell and Petter, 1980;Savage and Russell, 1983;Qiu, 1987;Kurtén and Garevski, 1989;Werdelin and Solounias, 1991;Koufos, 1992;Turner, 2001;Qiu et al., 2004;Turner et al., 2008;Liu and Qiu, 2009;Petrucci et al., 2013). However, Alba et al. (2015) recently demonstrated convincingly that authorship accreditation to Aymard is Fig. 4. Logarithmic ratio diagram comparing the teeth of Pachycrocuta brevirostris, Pachycrocuta perrieri and Parahyaena brunnea. ...
... Later, the view of Kurtén (1972), i.e., a single species without subspecies differentiation appearing in Eurasia, was widely accepted and supported by European scholars (e.g. Werdelin andSolounias, 1991, 1996;Turner and Antón, 1996;Werdelin, 1999;Turner et al., 2008;Dennell et al., 2008;Palmqvist et al., 2011). ...
Article
The giant short-faced hyena, Pachycrocuta brevirostris, was the largest and most spectacular bone-cracking hyena that ever existed. Although already extinct long ago, it was extensively distributed across Early and Middle Pleistocene localities throughout Eurasia. P. brevirostris is remarkable for a series of iconic craniodental adaptations in sway with a commitment to scavenging off carcasses of other large herbivores, such as the powerfully built skull and massive premolars, yet the most eye-catching aspect is the enormous size as emphasized by its vernacular name. The largest hitherto known skull of this species is the holotype from Sainzelles in Auvergne, France, discovered over a century ago. We hereby report a new discovery from the Jinyuan Cave in Dalian, China, which slightly surpasses the holotype in craniodental dimension, posing a new record in Pachycrocuta body size. Three skull fragments of the same individual unearthed from the site are described in detail herein and compared with the giant hyenas from both Europe and Asia. Thorough comparative studies show that the newly discovered specimens are very close to the holotype from Sainzelles both metrically and morphologically, which fully indicates the same morphotype as Europe, namely Pachycrocuta brevirostris brevirostris, surprisingly appeared in Northeast Asia as well, and further challenges the conventional concept of the population or subspecies. The giant hyenas from Asia were originally proposed as independent species, such as licenti or sinensis etc., but now broadly accepted as conspecifics of the European Pachycrocuta brevirostris, forming only different subspecies. The occurrence of P. b. brevirostris in Northeast Asia forcefully demonstrates that the differentiation is more of temporal than geographical significance, i.e., the subspecies of Eurasia basically constitute an ancestor-descendant sequence or clade evolved gradually from licenti to sinensis via brevirostris. Moreover, represented by the newly discovered specimens, the forerunner of P. b. brevirostris with transitional features appeared in Northeast Asia as early as 2.0 Ma at least, which implies the population of Europe most likely originated and dispersed from Northeast Asia. A hypothesis of “out of Northeast Asia” is hereby proposed for P. b. brevirostris. In present paper, a series of diagnostic craniodental characters, such as the carnassial, the accessory cuspids of premolar and the skull profile etc., are also discussed and re-valuated to investigate their potential significance in phylogenetic analysis.
... 4.2 Ma). Hyenas are of significant paleontological interest not only because they are frequently found in fossil assemblages of the Old World since the Miocene but also because (particularly since the Lower Pleistocene) many of these assemblages have been accumulated by them (Turner, Antón & Werdelin, 2008;Palmqvist et al., 2011). ...
... In agreement with Turner, Antón & Werdelin (2008), any investigations must operate within a clearly taxonomic framework. Although the taxonomic work on fossil hyenas has led to several important revisions (e.g., Howell & Petter, 1980;Kurtén & Werdelin, 1988;Werdelin, 1988aWerdelin, , 1988bSemenov, 2008;Tseng, Li & Wang, 2013), the seminal study by Werdelin & Solounias (1991) represents the most comprehensive framework for this family. ...
... Hyenids are mainly known by their durophagous members, but durophagy is not exclusive to this family, nor were all hyenids bone crackers ( Van Valkenburgh, 2007;Figueirido, Tseng & Martín-Serra, 2013). In spite of this family comprising more than 20 genera, practically all of them were assigned by Werdelin & Solounias (1996) to one of six ecomorphologies, which resemble living groups, namely: (1) civet-like insectivores/ omnivores, (2) mongoose-like insectivores/omnivores, (3) jackal-and wolf-like meat and bone eaters, (4) cursorial meat and bone eaters, (5) transitional bone crackers and (6) fully developed bone crackers (Werdelin & Solounias, 1996;Turner, Antón & Werdelin, 2008). These ecomorphological groups or ecomorphs are based on qualitative traits and can be seen as groups of genera sharing a more or less similar functional guild (or adaptive zone sensu Van Valen, 1971) by morphological analogy with living groups. ...
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We analyze the multivariate pattern of lower and upper cheek dentition for the family Hyaenidae along its evolutionary history. A total of 11,698 individual measurements of lengths and widths for the main postcanine teeth were collected for 54 extinct and three extant species of this family and analyzed by means of principal component analyses. Our results indicate that the functional aspects are better reflected by lower cheek dentition as a result of mosaic evolution. The multivariate structure captured by the three first principal components correspond to different adaptive strategies. The two first components characterize the main groups of ecomorphs, while hunting species separate from scavengers along the third axis. In the context of Hyaenidae, the post-canine cheek dentition of Parahyaena brunnea and Hyaena hyaena shows an extreme degree of specialization in scavenging.
... During the Late Pleistocene cave hyenas were the main non-anthropic accumulators of bone remains in caves (Fosse, 1997;Diedrich, 2014). As a consequence of its activity, hundreds of accumulations have been documented in European caves (Fosse, 1997;Fosse et al., 1998;Diedrich and Žák, 2006;Turner et al., 2008;Fourvel, 2012;Diedrich, 2014;Fourvel et al., 2014). In the Late Pleistocene of the Iberian Peninsula, the cave hyena was a common element in large mammal assemblages, being recorded from numerous sites Varela, 2011). ...
... The cave hyena enjoyed notable success, becoming a ubiquitous taxon in European faunal associations of the Late Pleistocene from the Urals to the Iberian Peninsula (Figure 15, Appendix 7) Carrión et al., 2006, Stuart andLister, 2014). Some authors hold that there are anatomical differences that are not sufficient for making a taxonomic distinction between individuals from the Eurasian Pleistocene and those from Africa (Turner et al., 2008). Results of studies of ancient DNA of these fossil remains suggest that the populations of European spotted hyena are a subset of the African population and not a distinct species (Rohland et al., 2005). ...
... In the last years, actuo-taphonomic studies show that Crocuta crocuta seldom creates large accumulations of bones despite having the ability. On the contrary, it is thought that the cave hyena made large accumulations of bones during the Late Pleistocene throughout Europe (Turner et al., 2008;see references in Diedrich, 2014;Fosse, 1997;Fourvel, 2012, Fourvel et al., 2015. Therefore, apart from the morphological differences between C. spelaea and C. crocuta, ethological differences might exist as well. ...
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A new Pleistocene paleontological site, Los Aprendices, located in the northwestern part of the Iberian Peninsula in the area of the Moncayo (Zaragoza) is presented. The layer with fossil remains has been dated by amino acid racemization to 143.8 ± 38.9 ka (earliest Late Pleistocene or latest Middle Pleistocene). Five mammal species have been identified in the assemblage: Crocuta spelaea (Goldfuss, 1823) Capra pyre-naica (Schinz, 1838), Lagomorpha indet, Arvicolidae indet and Galemys pyrenaicus (Geoffroy, 1811). The remains of C. spelaea represent a mostly complete skeleton in anatomical semi-connection. The hyena specimen represents the most complete skeleton ever recovered in Iberia and one of the most complete remains in Europe. It has been compared anatomically and biometrically with both European cave hyenas and extant spotted hyenas. In addition, a taphonomic study has been carried out in order to understand the origin and preservation of these exceptional remains. The results suggest rapid burial with few scavenging modifications putatively produced by a medium size carnivore. A review of the Pleistocene Iberian record of Crocuta spp. has been carried out, enabling us to establish one of the earliest records of C. spelaea in the recently discovered Los Aprendices cave, and also showing that the most extensive geographical distribution of this species occurred during the Late Pleistocene (MIS4-2).
... The more recently accepted species within Lycyaena include L. chaeretis, L. macrostoma, L. dubia, and L. crusafonti. Werdelin and Solounias (1991) suggest that L. chaeretis may be synonymous with L. dubia, although Turner et al. (2008) mention that would result in the former becoming the senior synonym of the latter, even though L. dubia is the more completely known of the taxa. Werdelin and Solounias (1991) noted only a small difference (M1 slightly more reduced in L. dubia). ...
... As a paraphyletic group (e.g. Werdelin and Solounias 1991;Semenov 2008;Turner et al. 2008), Semenov (2008 considered the 'ictitheres' to be comprised of two main groups, the Ictitheriinae sensu stricto (i.e. the true ictitheres) (e.g. Semenov 1989) and the Hyaenotheriini. ...
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Recently collected carnivoran specimens from the Siwalik Group of Pakistan are described and discussed. These specimens add to our knowledge of the previously known taxa while also adding to our biogeo-graphic, stratigraphic, and temporal knowledge of the carnivorans from the Siwaliks. At least eight distinct taxa are identified, and although most specimens are fragmentary, some hyaenid specimens are identified further (e.g. Lycyaena and Ictitherium). We identify and describe the first herpestid fossils from the Chinji Formation, the first hyaenid (Lycyaena cf. L. dubia) from the Tatrot Formation, and the first definitive occurrence of Ictitherium (Ictitherium cf. I. viverrinum) from the Dhok Pathan Formation. We report the first occurrences of several taxa from various sites in the Siwaliks of Pakistan, including the first reports of any carnivorans from Dhok Milan, Kohtehra, and Lawa. Individual sites show a wide range of carnivoran biodiversity, however larger scale trends are more discernible when comparing those between formations in the Siwaliks. Several taxa disappear over time from the carnivoran fauna of the Siwaliks, starting with barbourofelines, and continuing with amphicyonids (and non-carnivoran hyaenodontids). Felids and mus-telids are the most diverse carnivorans in the Siwaliks, while caniforms become more prevalent through time. ARTICLE HISTORY
... Ma; Sotnikova & Rook 2010). The niches of the two hyaenids, Pliocrocuta and Chasmaporthetes, are clearly separated (Figs 2-4) in almost all the examined parameters (except of the similar body size), reflecting thus a different mode of prey acquisition and exploitation: Pliocrocuta had marked bone-cracking abilities in its dentition indicative of a more scavenging behaviour, whereas Chasmaporthetes lunensis exploited its own hunted prey and was less adapted for cracking bones (Ferretti 1999;Antón et al. 2006;Turner et al. 2008;Tseng et al. 2011). Yet there exists partial niche overlap between Chasmaporthetes and the slightly larger (but still within the same body mass class) Acinonyx, as both were hypercarnivorous and pursuit predators (Fig., 4). ...
... However, cranial features and modern analogues are indicative for a solitary behaviour of the cheetah, which may have mainly focused on 50 kg class of prey (Hemmer et al. 2011). On the other hand, the pack-hunting tactic possibly employed by the hyaenid (Antón et al. 2006;Turner et al. 2008) may have allowed it to kill prey much larger than its own size and larger than what a single individual would succeed. Along with the incorporation of bones in the diet (Werdelin & Solounias 1996), reflecting a full exploitation of its own prey and/or opportunistic scavenging to a certain degree (Tseng et al. 2011), such differences may have reduced trophic conflicts. ...
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This study investigates the community structure, dynamics and evolution of the guilds of large carnivorans during the Pleistocene of Europe. Emphasis is given to important renewals, the composition of the guilds in terms of dietary preferences and foraging strategies, and to intraguild competition for access to food resources. For this purpose, cluster, principal component, and guild structure analyses are performed combining four ecological/behavioural parameters —body mass, diet, prey acquisition strategy, sociality— of large carnivorans that practice hunting and/or scavenging on large prey. Results show only minimal niche overlap, indicating that large predators may have reduced/avoided competition by almost exclusively occupying different niches, i.e. they did not compete for the same resources and/or employed different foraging strategy. Such niche partitioning and competition avoidance may have reduced the occurrences of potential trophic conflict and could explain the cooccurrence of a high diversity of large predators within the same broad feeding guild. Furthermore, the major predator guild remodeling took place close to the Early/Middle Pleistocene transition, when previously dominant carnivorans (e.g. Pachycrocuta, Megantereon, Acinonyx, Panthera gombaszoegensis) went extinct and new immigrants arrived (e.g. Panthera spelaea, Panthera pardus, Crocuta crocuta), forming the Galerian to Late Pleistocene guilds. Finally, the inclusion of the meat-eating Homo in the carnivore guild is discussed, including its possible impact to the demise of carnivoran diversity and accordingly of the several large carnivoran niches towards the end of the Pleistocene.
... Although Plioviverrops is among the earliest hyaenids to appear in the fossil record, as its earliest occurrences are in the MN4-5 (ca. 16.5 Ma) (Turner et al. 2008), Plioviverrops faventinus is the youngest species of this mongoose-like genus to appear in the fossil record ( Figure 7). The earliest records of Plioviverrops are scanty and not well characterised (Turner et al. 2008). ...
... 16.5 Ma) (Turner et al. 2008), Plioviverrops faventinus is the youngest species of this mongoose-like genus to appear in the fossil record ( Figure 7). The earliest records of Plioviverrops are scanty and not well characterised (Turner et al. 2008). Plioviverrops orbignyi from numerous localities of Greece dated between 9.7 and 7.0 Ma (Koufos 2011); and P. guerini from Spanish and Greece MN11-12 (8.7-7.0 ...
Article
Among the vertebrates found at Cava Monticino, carnivorans are by far the most abundant of all the large mammals. Five different taxa were recovered: one felid, two hyaenids, one canid, and one mustelid. The small-sized felid remains can be attributed to Felis christoli and seems to represent one of the earliest records of a true member of the genus Felis in Western Europe. Hyaenids at Cava Monticino are represented by the large wolf-sized and cursorial Lycyaena cf. chaeretis, and by the peculiar small Plioviverrops faventinus, the most abundant taxon of all. The latter is one of the most derived species of the genus and the last to appear in the fossil record of these mongoose-like hyaenids. The medium-sized canid recorded at Cava Monticino, Eucyon monticinensis, represent one of the oldest, certain record in the Old World of the genus Eucyon. It was a mesocarnivorous species that preyed on small vertebrates (abundantly recorded in the area of Cava Monticino during the Late Miocene). Lastly, mustelids are represented by the large relative of the extant honey badger, Mellivora benfieldi, whose record at Cava Monticino represents the northernmost record of the species and, presently, the only record of the genus outside of Africa.
... The dispersal of P. brevirostris in Europe would in all probability be from Asia, as this species was extinct in East Africa by these times (Werdelin, 1999), and its latest appearances are in the late Early Pleistocene (1.0-0.8 Ma; e.g., Sü ssenborn, Stranska Skala, Untermassfeld, Vallonnet, Cueva Victoria, Slivia, Incarcal, and Vallparadís) (Turner and Antó n, 1996;Arribas and Palmqvist, 1999;Turner et al., 2008;Palmqvist et al., 2011;Madurell-Malapeira et al., 2017). ...
... According to the information discussed above, the claim of Agustí and Lordkipanidze (2019) that ''the hyaenas from Dmanisi belong to be typical European Pliocene form, Pachycrocuta perrieri, and not the large hyaena Pachycrocuta brevirostris'' is incorrect: first, Pl. perrieri is the type species of the genus Pliocrocuta, not Pachycrocuta; second, the chronological range of Pl. perrieri in Europe spans from 4.2 to 2.0 Ma (MN15 to MN17 biozones; Turner et al., 2008;Palmqvist et al., 2011;Vinuesa et al., 2015), which means that the replacement of Pl. perrieri by P. brevirostris took place by the end of the Middle Villafranchian, prior to the age of those localities with typical Late Villafranchian faunas, like Dmanisi and Venta Micena. ...
Article
The archaeopaleontological site of Dmanisi in Georgia, dated to ∼1.8 Ma, provides evidence on the first hominin dispersal out of Africa, while the sites of Barranco León and Fuente Nueva-3 in Spain, dated to ∼1.4 Ma, record the earliest hominin settlements in Europe. However, a number of issues related to the dispersal route, the climatic conditions and the ecological scenario of this dispersal event are subject to debate. In a recent paper in L’anthropologie, Agustí and Lordkipanidze (2019) proposed an alternative scenario for the arrival of hominins in the Caucasus, which they conceived as a forest refugium area during the Early Pleistocene, and discarded that their dispersal coincided with that of other members of the Ethiopian and Asian faunas, like the sabertooth Megantereon whitei or the giant hyena Pachycrocuta brevirostris. Our review of these issues suggests that: (i) the elongated sabers and reduced postcanine teeth of African M. whitei limited the ability of this predator to process the prey carcass, which resulted in scavengeable resources for the Dmanisi hominins; (ii) the mass estimate in excess of 100 kg obtained for the trochlear perimeter of the distal humerus of the hyena from Dmanisi shows that it can be confidently ascribed to the genus Pachycrocuta; (iii) the postcranial anatomy of the Dmanisi hominins was not advantageous for scavenging tree-stored prey; (iv) the laterally flattened upper canines of M. whitei could not withstand the loads that would result from climbing a prey carcass into a tree; (v) paleobotanical analyses suggest a temperate grassland ecosystem in Dmanisi, not dominant forest conditions, with enhanced aridity in the level of hominin occupation; (vi) similarly, the low frequency of arboreal pollen in the Levantine Corridor at ∼1.8 Ma points to more arid conditions than today in this area; (vii) many archaeopaleontological sites of the Rift Valley and its extension towards the Red Sea, the Levant and the Caucasus show evidence of tectonic, volcanic and/or hydrothermal events; and (viii) the delay of 400 ka in the arrival of hominins in Western Europe did not result from a lower availability of scavengeable resources.
... Two bone-cracking hyaenids are reported in the European Early Pleistocene: Pliocrocuta perrieri (late Pliocene-Early Pleistocene; about 4.2-2.0 Ma) (Turner et al. 2008;Vinuesa et al. 2014) and Pachycrocuta breviro-stris (Early-Middle Pleistocene; about 2.0-0.8 Ma) (Turner et al. 2008;Madurell-Malapeira et al. 2010). ...
... Ma) (Turner et al. 2008;Vinuesa et al. 2014) and Pachycrocuta breviro-stris (Early-Middle Pleistocene; about 2.0-0.8 Ma) (Turner et al. 2008;Madurell-Malapeira et al. 2010). Unfortunately, it is not possible to assign the PSL coprolites to one or the other species on the basis of morphology alone. ...
Article
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Most of the research on fossil mammals from Umbria (central Italy) has been carried out in the southwestern branch of the Tiber basin, due to its paleontological richness. This portion of the basin extends from Perugia to Terni and corresponds to a well-defined half-graben filled with fluvial-lacustrine deposits. The paleontological sample presented here was discovered in a sand and gravel quarry at Podere San Lorenzo, East of the town of Deruta. The stratigraphic succession exposed in the quarry is no longer visible, but we describe here a new outcrop (Palazzone), which is not far from Podere San Lorenzo and shows comparable facies associations. The two successions were deposited in a fluvial environment characterized by an average reduction of the hydrodynamic energy from the bottom upwards. They are referred to the Early Pleistocene Santa Maria di Ciciliano Subsyntheme (Madonna dei Bagni Lithofacies). Large mammal remains are attributed to Mammuthus cf. meridionalis (Nesti, 1825), Stephanorhinus etruscus (Falconer, 1859), Equus stenonis Cocchi, 1867, Leptobos cf. etruscus (Falconer, 1868), ‘Pseudodama’ nestii (Azzaroli, 1947), and Sus strozzii Forsyth Major, 1881. Some hyena coprolites are also reported. The assemblage is typical of the early Late Villafranchian Land Mammal Age and can be referred to the Olivola/Tasso Faunal Units (about 2.0–1.8 Ma). This is in agreement with the alleged age of some other assemblages found in the southwestern branch of the Tiber basin (e.g., Torre Picchio, Villa San Faustino, Colle Sant’Andrea, Pantalla).
... 9.6-4.9 Ma (Turner et al. 2008). It is recorded from several Eurasian localities, mainly on the basis of fragmentary material. ...
... This similarity between A. eximia and Proteles suggests that the former displays a more primitive endocranial morphology than its extant ecomorphological analogs. Given that A. eximia is customarily related to the extant spotted hyena (Crocuta crocuta) based on external craniodental characters (Turner et al. 2008), the observed endocranial similarities with Proteles are probably indicative of a less elaborated social behavior compared to extant bone-cracking hyenas. The preliminary work reported here represents a first step for the study of the endocranial morphology of A. eximia. ...
... During the MPR, a faunal replacement took place in Europe with the appearance of new carnivores (see below), an increase in herbivore richness and a change in the structure of the mammalian communities (Turner 1992;Palombo 2010Palombo , 2014Rodríguez et al. 2012). Arribas and Palmqvist (1999) and Turner et al. (2008) argued that the sabretooth cats M. whitei and H. latidens were replaced during the Middle Pleistocene by the modern pantherine felids, which exploited their prey in more depth, and that this resulted in the loss of a regular source of scavengeable carcasses for both the hominins and the giant hyaenas (however, note that Homotherium survived in NW Europe until the Late Pleistocene; Reumer et al. 2003). Under these new ecological circumstances, the trophic niche that hominins exploited vanished, and this forced them towards behavioural improvements that resulted in the development of the more effective Acheulean handaxes, which replaced the technologically less elaborated Oldowan flakes of the earlier populations. ...
Article
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The late Early Pleistocene archaeological site of Fuente Nueva 3 (Orce, Guadix-Baza Depression, SE Spain), dated to ~1.4 Ma, provides evidence on the subsistence strategies of the first hominin population that dispersed in Western Europe. The site preserves Oldowan tool assemblages associated with abundant remains of large mammals. A small proportion of these remains show cut marks and percussion marks resulting from defleshing and bone fracturing, and a small proportion of bones also show tooth marks. Previous taphonomic studies of FN3 suggested that the hominins had secondary access to the prey leftovers abandoned by sabretooth cats and other primary predators. However, a recent analysis by Yravedra et al. (2021) of the frequency of anthropogenic marks and tooth marks has concluded that the hominins had primary access to the carcasses of a wide variety of ungulate prey, even though the frequency of evisceration marks is strikingly low. In this rebuttal, we analyse the patterns of bone preservation in FN3, which show that the exploitation of bone marrow by the hominins after hammerstone breakage was a usual activity at the site. Our study also reviews the evidence available on the lesser abilities of sabretooth cats for carcass processing compared to pantherine felids. This reinforces the hypothesis that primary predators provided the hominins the opportunity to scavenge sizeable chunks of meat and bone marrow of their prey carcasses before the arrival of hyaenas. Finally, we also provide new inferences on resource availability and competition intensity among the members of the carnivore guild in FN3, which reinforce our interpretation that a secondary access by the Oldowan hominins to the prey leftovers of sabretooth cats was an optimal foraging strategy in the Guadix-Baza Depression.
... Their interaction is reflected in the exploitation of prey by hominins and carnivores and is commonly studied using the taphonomic analysis of post-cranial and dental elements (Marciszak 2012). C. spelaea shared the environment with hominins in Europe for millennia (Turner et al. 2008), and they even followed similar migratory routes with comparable timing patterns in their dispersal from Africa (Kolendrianou et al. 2020). In the case of Palaeolithic caves in the Swabian Jura Caves (Germany), Discamps et al. (2012) suggested an increase of small carnivores over time, and Camarós et al. (2016) agreed that there is a clear decrease in the size of the carnivores that were active from the Middle to the Upper Palaeolithic. ...
Article
Mani peninsula is considered an important region for the Palaeolithic research of Greece. A considerable number of sites have been excavated to date, containing deposits with cultural remains from the Middle Palaeolithic, the Upper Palaeolithic, and the Neolithic period. Melitzia Cave is a cave locality filled with Upper Pleistocene deposits, yielding in the upper stratigraphic units fossil remains of numerous large mammals and small vertebrates along with cultural remains, indicating that the cave was inhabited by both humans and carnivores. The layers date to the Upper Pleistocene, and more specifically between 46,448–44,553 BP and 11,150–10,680 BP. The present study focuses on the recovered carnivore assemblage of Melitzia Cave, which has revealed the presence of Crocuta spelaea, Vulpes vulpes, Felis silvestris, Meles meles, Mustela nivalis, Martes sp., ?Ursus sp., and Carnivora indet., described in detail herein. This species composition is typical for the late Pleistocene of Southeastern Europe.
... Hyenas were once an ecologically diverse group that showed adaptations to many different feeding behaviors, from the probable insectivory/omnivory of civetlike and mongoose-like species, to hypercarnivory and "bone-cracking" (Kurtén, 1956;Howell & Petter, 1980;Werdelin & Solounias, 1991Semenov, 2008;Turner et al., 2008;Tseng et al., 2013;Coca-Ortega & Pérez-Claros, 2019;Koufos, 2021;Iurino et al., 2022;Lewis & Werdelin, 2022). The latter is almost the only ecomorphotype (or ecomorph; Werdelin & Solounias, 1996) that survived to the present day. ...
Article
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We redescribe and revise the taxonomic attribution of a lost hyena mandible recovered from Paciano (Umbria, Italy), originally reported in the early 1900s, by comparing it with relevant samples of Pliocene, Pleistocene, and extant species. The mandible of the Paciano hyena was initially attributed to Hyaena striata (= Hyaena hyaena) and subsequently listed as a record of the giant hyena Pachycrocuta brevirostris, but is here assigned to another “bone-cracking” hyena, the Pliocene-Early Pleistocene Pliocrocuta perrieri. The Paciano hyena contributes to the discussion on the relationships and turnover between Pl. perrieri and P. brevirostris. On the one hand, the two species are very similar in craniodental morphology; their isolated remains are often separated by size; and P. brevirostris is thought to derive from a large-sized population of Pl. perrieri. On the other, a larger size is not an aspect to disregard in hyenas as it usually correlates with the acquisition or better development of “bone-cracking” features; remains attesting the co-occurrence of Pl. perrieri and P. brevirostris are known from some sites dated at around 2.0–1.8 Ma; and the observed size differences between the two species exceed those recorded between extant and (at times) sympatric species. Therefore, taken alone the competition with P. brevirostris does not explain the extinction of Pl. perrieri, but considering it together with the concurrent spread and resulting peak of carnivoran diversity in Late Villafranchian faunas might. Pliocrocuta perrieri was outcompeted by its larger descendent in scavenging carcasses, while other carnivorans limited its options to deviate to other resources or adopt a different feeding behavior (e.g., active hunting). The case of Paciano is also significant in that it offers an example of the importance and feasibility to reconsider historical collections, even when the original material is lost entirely. In general, the seek for “novelty” that permeates current scientific literature ensues in few descriptions or reconsiderations of historical samples, especially if the new examinations confirm old results, but we emphasize the significance of such efforts in making old data truly available for the scientific community. In addition, the hyena from Paciano has a biochronological and stratigraphic value, confirming the occurrence in western Umbria of a depositional cycle older than the late Early Pleistocene one (well represented by the rich assemblages from Pietrafitta and Selvella; Farneta Faunal Unit).
... Sometimes it is difficult to distinguish them from each other. As a result, P. pyrenaica was even questioned to be a probable synonym of P. perrieri (Werdelin and Solounias, 1991;Turner et al., 2008). The Yegou specimen V 18835 bears the typical morphology of P. pyrenaica (based on the standard of Howell and Petter, 1980), which provides powerful surport for the validity of the species. ...
Article
Full-text available
Currently, there are still different views regarding the chronology of the Late Cenozoic deposits in the Nihewan Basin, which results from the contradiction between biostratigraphic correlations based on mammalian fossils and magnetostratigraphic dating results. Biostratigraphic correlations indicate that the aeolian red clay exposed in the Sanggan River canyon, the fluvio-lacustrine red clay with sands and gravels, and the sandy clay of swamp facies on both sides of the lower reaches of the Huliu River belong to the Upper Pliocene, whereas the magnetostratigraphic dating usually correlates them to the Lower Pleistocene. In October 2011, a collection of mammalian fossils was unearthed from a block of collapsed deposits at Yegou in the Nihewan Basin, which is about 300 m north of the Laowogou section that is well known for the Pliocene mammalian fossils from its lower part. The Yegou fossils are identified herein as 10 species in 9 genera: Nyctereutes tingi, N. sinensis, Pachycrocuta pyrenaica, Homotherium sp., Hipparion (Plesiohipparion) houfenense, Dicerorhinus sp., Muntiacus sp., Axis shansius, Gazella blacki, and Paracamelus sp. The fauna is quite different from the classic Early Pleistocene Nihewan Fauna in composition and provides new evidence for the existence of the Upper Pliocene in the Nihewan Basin. Based on a systematic description of the fauna, its composition and geological age are discussed, and the compositional features of large mammals of the Late Pliocene and the Early Pleistocene mammalian faunas in the Nihewan Basin are summarized.
... In Europe, five Late Miocene species Hyaenictitherium hyaenoides (Zdansky 1924), H. wongii (Zdansky 1924), H. intuberculatum (Ozansoy, 1965), H. parvum (Khomenko, 1914) and H. venator (Semenov, 1989) have been described while from Africa, two Late Miocene species; H. minimum and Hyaenictitherium cf. H. parvum (Werdelin 2003), have been reported, followed by H. namaquensis in the Early Pliocene of Libya, South Africa, and Kenya (Werdelin and Solounias 1991;Werdelin et al. 1994;Morales et al. 2005;Turner et al. 2008). The genus greatly expanded its geographic range in Eurasia between MN11 and MN13 (and equivalent ages outside of Europe), reaching China to the east and the Indian subcontinent to the south during the Late Miocene. ...
Article
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New dental material-one maxilla bearing P3 and P4-of Early Pliocene hyaenid Hyaenictitherium pilgrimi from the Dhok Pathan Formation of Hasnot is described and discussed. The presence of this hyaenid from the Early Pliocene (Ruscinian equivalent) deposits at Hasnot is an important fossil record of this species from the Siwalik continental deposits of Pakistan. The purpose of the paper is to provide more information about the fossil record and its stratigraphic extension from Late Miocene to Early Pliocene from the Siwalik continental deposits.
... In Europe, five Late Miocene species Hyaenictitherium hyaenoides (Zdansky 1924), H. wongii (Zdansky 1924), H. intuberculatum (Ozansoy, 1965), H. parvum (Khomenko, 1914) and H. venator (Semenov, 1989) have been described while from Africa, two Late Miocene species; H. minimum and Hyaenictitherium cf. H. parvum (Werdelin 2003), have been reported, followed by H. namaquensis in the Early Pliocene of Libya, South Africa, and Kenya (Werdelin and Solounias 1991;Werdelin et al. 1994;Morales et al. 2005;Turner et al. 2008). The genus greatly expanded its geographic range in Eurasia between MN11 and MN13 (and equivalent ages outside of Europe), reaching China to the east and the Indian subcontinent to the south during the Late Miocene. ...
... Hyenas belong to the order Carnivora, suborder of Feliformia and the family of Hyanidae. 1 Although phylogenetically closer to cats, hyenas are more similar to dogs in behaviour and body composition because both dogs and hyenas are runners who catch prey with their teeth rather than claws and eat quickly. 2 There have been several reports of spotted hyenas being infected with the canine distemper virus (CDV). [3][4][5][6] Haas et al. 4 reported the prevalence of distemper disease in a group of spotted hyenas. ...
Article
A 6‐month‐old free‐living female striped hyena (Hyaena hyaena) was referred to the Veterinary Teaching Hospital of Ferdowsi University of Mashhad, Mashhad, Iran with lethargy and seizures. The general and neurological examinations revealed normal heart rate, polypnoea, abnormal respiratory sounds, decreased consciousness and no postural reactions in limbs. The animal had generalised seizures every 20 minutes lasting about 1 minute. To control seizures, 10 mg diazepam (0.7 mg/kg) was injected intravenously four times every 2 minutes. With this treatment, the animal's seizures stopped. Based on characteristic clinical signs, lymphopaenia, and positive rapid kit and RT‐PCR results, canine distemper virus (CDV) was confirmed. The animal developed general seizures again 12 hours after referral and died during the seizures. This is the first case report of status epilepticus due to CDV in a striped hyena that did not respond to anticonvulsant therapies and caused the death of the animal.
... With the rapid uplift of the QTP, the environment changed into a dry and cold climate (Lu and Guo, 2013), and species differentiation of Cyclophyllidea was accelerated by the rapid adaptive evolution of their hosts and geographical isolation caused by the radiation of hosts to the Palaearctic (Favre et al., 2015;Xing and Ree, 2017;Päckert et al., 2020). Finally, in the last 2 million years, Cyclophyllidea differentiation demonstrated an accelerated diversification based on cox1 + nad1 divergence tree (Figure 5), which may be related to the evolution and broad distribution of mammals in Eurasia and the associated population expansion and migration of hominids from Africa to Asia (Dennell, 2004;Rohland et al., 2005;Brugal and Croitor, 2007;Turner et al., 2008;Klein, 2009;Terefe et al., 2014;Wang et al., 2016). The Host-parasite Database of Natural History Museum (HPDNHM) found that most tapeworm were prevalent mainly in the Palaearctic, 5 which is also consistent with the viewpoint that the order Cyclophyllidea originated from the QTP. ...
Article
Full-text available
The Cyclophyllidea comprises the most species-rich order of tapeworms (Platyhelminthes, Cestoda) and includes species with some of the most severe health impact on wildlife, livestock, and humans. We collected seven Cyclophyllidea specimens from rodents in Qinghai-Tibet Plateau (QTP) and its surrounding mountain systems, of which four specimens in QTP were unsequenced, representing “putative new species.” Their complete mitochondrial ( mt ) genomes were sequenced and annotated. Phylogenetic reconstruction of partial 28S rDNA, cox 1 and nad 1 datasets provided high bootstrap frequency support for the categorization of three “putative new species,” assigning each, respectively, to the genera Mesocestoides , Paranoplocephala , and Mosgovoyia , and revealing that some species and families in these three datasets, which contain 291 species from nine families, may require taxonomic revision. The partial 18S rDNA phylogeny of 29 species from Taeniidae provided high bootstrap frequency support for the categorization of the “putative new species” in the genus Hydatigera . Combined with the current investigation, the other three known Taeniidae species found in this study were Taenia caixuepengi , T. crassiceps , and Versteria mustelae and may be widely distributed in western China. Estimates of divergence time based on cox 1 + nad 1 fragment and mt protein-coding genes (PCGs) showed that the differentiation rate of Cyclophyllidea species was strongly associated with the rate of change in the biogeographic scenarios, likely caused by the uplift of the QTP; i.e., species differentiation of Cyclophyllidea might be driven by host-parasite co-evolution caused by the uplift of QTP. We propose an “out of QTP” hypothesis for the radiation of these cyclophyllidean tapeworms.
... The species arrived in Europe during the early Pleistocene (1.99 Ma, Olivola Faunal Unit; Napoleone et al. 2003), replaces Pliocrocuta perrieri and Chasmaporthetes lunensis, and extincted at the end of early Pleistocene (~1.0-0.8 Ma) (Turner et al. 2008). Its extinction is related to that of Homotherium and Megantereon, the two main hunting predators of early Pleistocene, which provided a lot of carcases, the main food for Pachycrocuta (Turner and Anton 1996). ...
Article
Full-text available
The mammal fauna from Libakos was collected by H. Eltgen (Technical University of Clausthal, Germany) during the end of the 1960s and studied by K. Steensma. The locality is situated within the fluviolacustrine-fluvioterrestrial deposits of the Grevena–Neapolis basin (Macedonia, Greece), which are considered as Pliocene-Pleistocene. The carnivoran collection of Libakos is poor and fragmentary with doubtful or limited determinations. The reconsideration of the carnivorans indicated the presence of the canids Caniscf. arnensis and Canis cf. etruscus, the mustelid Pannonictis nestii recognised for first time in Greece, the large-sized hyaenid Pachycrocuta brevirostris and the machairodontine Homotherium latidens. The available biochronological data, the stratigraphy and the comparison with other well-dated Villafranchian faunas of Greece, indicate a late Villafranchian age for Libakos fauna, more precisely between ~2.0 and 1.5 Ma.
... Sometimes it is difficult to distinguish them from each other. As a result, P. pyrenaica was even questioned to be a probable synonym of P. perrieri (Werdelin and Solounias, 1991;Turner et al., 2008). The Yegou specimen V 18835 bears the typical morphology of P. pyrenaica (based on the standard of Howell and Petter, 1980), which provides powerful surport for the validity of the species. ...
Preprint
Currently, there are still different views regarding the chronology of the Late Cenozoic deposits in the Nihewan Basin, which results from the contradiction between biostratigraphic correlations based on mammalian fossils and magnetostratigraphic dating results. Biostratigraphic correlations indicate that the aeolian red clay exposed in the Sanggan River canyon, the fluvio-lacustrine red clay with sands and gravels, and the sandy clay of swamp facies on both sides of the lower reaches of the Huliu River belong to the Upper Pliocene, whereas the magnetostratigraphic dating usually correlates them to the Lower Pleistocene. In October 2011, a collection of mammalian fossils was unearthed from a block of collapsed deposits at Yegou in the Nihewan Basin, which is about 300 m north of the Laowogou section which is well known for the Pliocene mammalian fossils from its lower part. The Yegou fossils are identified herein as 10 species in 9 genera: Nyctereutes tingi, N. sinensis, Pachycrocuta pyrenaica, Homotherium sp., Hipparion (Plesiohipparion) houfenense, Dicerorhinus sp., Muntiacus sp., Axis shansius, Gazella blacki, and Paracamelus sp. The fauna is quite different from the classic Early Pleistocene Nihewan Fauna in composition and provides new evidence for the existence of the Upper Pliocene in the Nihewan Basin. Based on a systematic description of the fauna, its composition and geological age are discussed, and the compositional features of large mammals of the Late Pliocene and the Early Pleistocene mammalian faunas in the Nihewan Basin are summarized.
... ★Belbus beaumonti (Qiu, 1987) Nomenclatural and Taxonomical History Hyaena sp. in Beaumont 1968 (initial identification); Hyaenictitherium hyaenoides in Howel and Petter 1980 (new combination); Hyaenictis beaumonti in Qiu 1987 (new species); Belbus beaumonti in Werdelin and Solounias 1991 (new genus and combination). The taxonomic history of this material is long and complicated, referred under various generic and specific names (Werdelin and Solounias 1991;Turner et al. 2008). ...
Chapter
The family Percrocutidae was erected to include some Miocene and Pliocene hyaena-like carnivoran taxa of Eurasia and Africa, which were included to the Hyaenidae for a long time. The percrocutids were dispersed from Spain to China and in the Eastern Mediterranean region are traced in Serbia, Bulgaria, and Turkey. In Greece, Percrocutidae are scarcely known from the middle and late Miocene of northern Greece and Samos Island. It is, however, quite diverse and the three identified certain species (two of them new) belong to the genera Percrocuta, Dinocrocuta, and Belbus. The members of the family are characterized by the absence of the M2/m2 and the enlarged premolars, mainly P3/p3.KeywordsCarnivoraPercrocutidae Belbus Dinocrocuta Percrocuta NeogeneMiocene
... The earliest representatives of Chasmaporthetes are from the late Miocene and are found in China, represented by C. exitelus and in Greece represented by C. bonisi. The precise taxonomic attribution of C. bonisi and the possible synonymy between these two taxa remains to be clarified, however (Werdelin and Solounias, 1991;Turner et al., 2008;Tseng et al., 2013;Vinuesa et al., 2016). The oldest uncontroversial species of the genus is C. australis, which occurs during the Messinian in Africa, where C. nitidula is also recorded during the Pliocene and early Pleistocene. ...
... Among the hyenids, Hyaena prisca has been considered so far as an ancestral form of the extant striped hyena Hyaena hyaena (Kurten, 1968;Werdelin & Solounias, 1991;Turner, 1990;Turner et al., 2008), although recent research suggests a potential phyletic relationship with the extant brown species Parahyaena brunnea (Arribas & Garrido, 2008;Brugal et al., 2012). Hyaena prisca is present in some Pleistocene sites especially in Southern Europe and seems more characteristic of the Middle Pleistocene (Kurten, 1968;Brugal et al., 2020). ...
Article
The archeo-paleontological site of Mas des Caves at Lunel-Viel (Hérault), in Southeastern France, is an important site, well-known for its diversified vertebrate remains dated to the second half of the Middle Pleistocene. Following a brief presentation of the historical context, preliminary data, collected through new research, are discussed, focusing on the general fossil distribution and new analyses of leporids, carnivores, rhinocerotids and cervids to better define their taxonomy, paleoecology and biochronology.
... The smallest family of Carnivora, Hyaenidae, includes only four extant species: spotted hyena (Crocuta crocuta), brown hyena (Parahyaena brunnea) and aardwolf (Proteles cristata) in Africa, and striped hyena (Hyaena hyaena) in both Africa and western Asia. However, this family formerly comprised over 80 species known from the fossil record and inhabited a wide geographical range across Eurasia, Africa and northern America [1,2]. Despite Hyaenidae having sustained an obvious decline in diversity and geographical distribution, extant hyenas still occupy a broad range of ecological niches, including the highly specialized insectivore niche occupied by the aardwolf and the more generalist bonecracking predatory and scavenging niche occupied by the other three extant species [3,4]. ...
Article
Cave hyenas (genus Crocuta) are extinct bone-cracking carnivores from the family Hyaenidae and are generally split into two taxa that correspond to a European/Eurasian and an (East) Asian lineage. They are close relatives of the extant African spotted hyenas, the only extant member of the genus Crocuta. Cave hyenas inhabited a wide range across Eurasia during the Pleistocene, but became extinct at the end of the Late Pleistocene. Using genetic and genomic datasets, previous studies have proposed different scenarios about the evolutionary history of Crocuta. However, causes of the extinction of cave hyenas are widely speculative and samples from China are severely understudied. In this study, we assembled near-complete mitochondrial genomes from two cave hyenas from northeastern China dating to 20 240 and 20 253 calBP, representing the youngest directly dated fossils of Crocuta in Asia. Phylogenetic analyses suggest a monophyletic clade of these two samples within a deeply diverging mitochondrial haplogroup of Crocuta. Bayesian analyses suggest that the split of this Asian cave hyena mitochondrial lineage from their European and African relatives occurred approximately 1.85 Ma (95% CI 1.62-2.09 Ma), which is broadly concordant with the earliest Eurasian Crocuta fossil dating to approximately 2 Ma. Comparisons of mean genetic distance indicate that cave hyenas harboured higher genetic diversity than extant spotted hyenas, brown hyenas and aardwolves, but this is probably at least partially due to the fact that their mitochondrial lineages do not represent a monophyletic group, although this is also true for extant spotted hyenas. Moreover, the joint female effective population size of Crocuta (both cave hyenas and extant spotted hyenas) has sustained two declines during the Late Pleistocene. Combining this mitochondrial phylogeny, previous nuclear findings and fossil records, we discuss the possible relationship of fossil Crocuta in China and the extinction of cave hyenas.
... Briefly, A. eximia was originally described as Hyaena eximia Roth and Wagner, 1854, then attributed to the genus Crocuta Kaup, 1829, and finally allocated to the genus Adcrocuta Kretzoi, 1938. Werdelin andSolounias (1991) have an extensive list of synonymies for A. eximia, which is monospecific with A. eximia being the only species within the genus Adcrocuta (Turner et al., 2008). It has also been argued that A. eximia can be compared to extant spotted hyena Crocuta crocuta (de Bonis, 2005). ...
... Members of the family Hyaenidae (part of Order Carnivora, which also includes wolves, cats, seals, weasels, and relatives) have an extensive fossil record across the Old World beginning in the early Miocene, with >60 extinct hyaenid taxa having been described. They are often the most abundant predators at fossil localities across Eurasia, and are considered to have played key ecological roles as mesopredators and scavengers (Turner et al., 2008). They were, however, absent from North America until dispersal (presumably via the Beringian land bridge) during the Blancan North American Land Mammal Age (NALMA, Figure S1), ~4 .7 Ma (mega-annum) ago. ...
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The northern region of Beringia is ecologically and biogeographically significant as a corridor for biotic dispersals between the Old and New Worlds. Large mammalian predators from Beringia are exceedingly rare in the fossil record, even though carnivore diversity in the past was much higher than it is in this region at present. Here we report the first fossils of cursorial hyenas, Chasmaporthetes, in Beringia and north of the Arctic Circle. Two isolated teeth recovered in the Old Crow Basin, Yukon Territory, Canada, were identified amongst over 50,000 known fossil mammal specimens recovered from over a century of collecting in the Old Crow Basin. These rare records fill an important intermediary locale in the more than 10,000 km geographic distance between previously known New and Old World records of this lineage. The Pleistocene age of these fossils, together with its Arctic Circle occurrence, necessitate a rethinking of the role of large-bodied hunter-scavengers in Ice Age megafaunas in North America, and the implications of lacking an important energy flow modifier in present day North American food webs.
... The Fort Ternan material includes two dp4 that both match the morphology of dp4 in Percrocutidae. In addition to these features, all of the Fort Ternan percrocutid specimens are considerably more derived than contemporaneous Eurasian Hyaenidae, which at that time (ca MN 6 or early MN 7/8 depending on opinion in the European mammal zonation) were confined to small, omnivorous forms such as Protictitherium and Plioviverrops (Turner et al. 2008): type 2 hyenas in the classification of Werdelin & Solounias (1996). ...
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Fort Ternan is a middle Miocene (c. 13.7-13.8 Ma) site famous for its fossils of Kenyapithecus wickeri Leakey, 1962, considered the earliest African hominoid. Herein, the Carnivora and Hyaenodonta from this site are described and placed in their temporal context, showing the middle Miocene to be a time of transition from archaic carnivores of the early Miocene and carnivores of more modern aspect from the late Miocene. Fort Ternan includes: Amphicyonidae represented by ?Myacyon peignei n. sp., a new form distinguished by its hypercarnivorous m1, P4 with large protocone shelf, and M1 with reduced lingual shelf; Barbourofelidae, represented by a derived form; Percrocutidae, represented by abundant material of Percrocuta tobieni Crusafont & Aguirre, 1971; Viverridae, represented by the paradoxurines Kanuites lewisae Dehghani & Werdelin, 2008, and cf. Orangictis Morales & Pickford, 2005, and a putative viverrine; and Hyaenodonta represented by the teratodontine Dissopsalis pyroclas-ticus Savage, 1965 and a very large hyainailourine. This assemblage is a melange of forms harkening back to the early Miocene (the Hyaenodonta and Amphicyonidae), an evolving, still extant lineage (Viverridae), and more typical late Miocene forms (derived Barbourofelidae and Percrocutidae). © 2019, Museum National d'Histoire Naturelle. All rights reserved.
... Classification rates are very high in all cases; however, Another point in favor of ML algorithms is their ability to separate the two types of studied carnivores. While the wolves studied present the problem of their captivity, as previously discussed by other authors (Sala et al. 2014;Gidna et al. 2015;Domínguez-Rodrigo et al. 2015), it is worth noting that, theoretically, hyenas and wolves are durophagous taxa that may be competitors for resources in certain ecosystems (Turner et al. 2008), and the way they break bones is not the same (Werdelin 1989;Ferretti 2007;Tseng and Binder 2010). Furthermore, in the case of hyenids, bone breakage is produced by the premolars; meanwhile, canids break bones with the first and/or the second set of molars (Tseng and Binder 2010). ...
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The analysis of bone breakage has always been underrepresented in taphonomic studies. Analysts, thus, lose the opportunity to resolve an important part of the equifinality related to activities that hominins and different types of carnivores may produce. Recent studies have shown that the use of powerful machine learning (ML) algorithms allow the accurate classification of bone surface modifications (BSM). Here, we present an experimental study, applying these algorithms to the analysis of bone breakage patterns. This statistical methodology allows the correct classification of three different assemblages which have been generated anthropogenically and by the activity of carnivores (i.e., hyenas and wolves). ML algorithms applied to a multivariate set of properties of broken bone specimens yielded an accuracy of 95% and were higher in classifying agency without the need to include information from BSM. This paper proposes a methodological approach that opens the door to improve our understanding of referential frameworks regarding bone breakage and to determine agency in prehistoric bone breakage processes.
... The family has a rich fossil history but is now restricted to only four extant species (Koepfli et al. 2006) . The abundance of fossil specimens has led to a number of studies focused on the taxonomic relationships within fossil Hyaenidae (Werdelin and Solounias 1991;Turner et al. 2008) . Little attention, however, has been focused towards the evolution and population status of the extant species, especially on a molecular level. ...
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The sequencing of the human genome in the early 2000s led to an increased interest in cheap and fast sequencing technologies. This interest culminated in the advent of next generation sequencing (NGS). A number of different NGS platforms have arisen since then all promising to do the same thing, i.e. produce large amounts of genetic information for relatively low costs compared to more traditional methods such as Sanger sequencing. The capabilities of NGS meant that researchers were no longer bound to species for which a lot of previous work had already been done (e.g. model organisms and humans) enabling a shift in research towards more novel and diverse species of interest. This capability has greatly benefitted many fields within the biological sciences, one of which being the field of evolutionary biology. Researchers have begun to move away from the study of laboratory model organisms to wild, natural populations and species which has greatly expanded our knowledge of evolution. NGS boasts a number of benefits over more traditional sequencing approaches. The main benefit comes from the capability to generate information for drastically more loci for a fraction of the cost. This is hugely beneficial to the study of wild animals as, even when large numbers of individuals are unobtainable, the amount of data produced still allows for accurate, reliable population and species level results from a small selection of individuals. The use of NGS to study species for which little to no previous research has been carried out on and the production of novel evolutionary information and reference datasets for the greater scientific community were the focuses of this thesis. Two studies in this thesis focused on producing novel mitochondrial genomes from shotgun sequencing data through iterative mapping, bypassing the need for a close relative to serve as a reference sequence. These mitochondrial genomes were then used to infer species level relationships through phylogenetic analyses. The first of these studies involved reconstructing a complete mitochondrial genome of the bat eared fox (Otocyon megalotis). Phylogenetic analyses of the mitochondrial genome confidently placed the bat eared fox as sister to the clade consisting of the raccoon dog and true foxes within the canidae family. The next study also involved reconstructing a mitochondrial genome but in this case from the extinct Macrauchenia of South America. As this study utilised ancient DNA, it involved a lot of parameter testing, quality controls and strict thresholds to obtain a near complete mitochondrial genome devoid of contamination known to plague ancient DNA studies. Phylogenetic analyses confidently placed Macrauchenia as sister to all living representatives of Perissodactyla with a divergence time of ~66 million years ago. The third and final study of this thesis involved de novo assemblies of both nuclear and mitochondrial genomes from brown and striped hyena and focussed on demographic, genetic diversity and population genomic analyses within the brown hyena. Previous studies of the brown hyena hinted at very low levels of genomic diversity and, perhaps due to this, were unable to find any notable population structure across its range. By incorporating a large number of genetic loci, in the form of complete nuclear genomes, population structure within the brown hyena was uncovered. On top of this, genomic diversity levels were compared to a number of other species. Results showed the brown hyena to have the lowest genomic diversity out of all species included in the study which was perhaps caused by a continuous and ongoing decline in effective population size that started about one million years ago and dramatically accelerated towards the end of the Pleistocene. The studies within this thesis show the power NGS sequencing has and its utility within evolutionary biology. The most notable capabilities outlined in this thesis involve the study of species for which no reference data is available and in the production of large amounts of data, providing evolutionary answers at the species and population level that data produced using more traditional techniques simply could not.
... Intraguild competition has also been claimed to be an important factor for hominin survival. For example, in the scenario described above, hominins would have had to confront the giant hyaena (Pachycrocuta brevirrostris), a superb competitor for carrion (Turner et al., 2008). Thus, analyzing primary production, resource availability and carnivoran paleoecology in Europe during the late Early Pleistocene and the beginning of the Middle Pleistocene is highly relevant to the hot debate over the effect of the environment on the dispersal of Homo out of Africa and, more specifically, the early settlement of Europe (Palombo, 2013;. ...
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Carrying capacity, the maximum biomass that an ecosystem can sustain over the long term, strongly influences several ecological processes and it is also one of the main determinants of biodiversity. Here, we estimate the carrying capacity (CC) of the late Early and early Middle Pleistocene ecosystems of Europe, using equations describing the relationship between CC and climatic variables observed in the present, as well as maps of inferred paleotemperature and paleoprecipitation. Maps of paleoclimate values were interpolated from the composite benthic stable oxygen isotope ratios and a transfer function was used to estimate ungulate carrying capacity (CCU) from the interpolated mean annual temperature and annual precipitation values. Carnivoran carrying capacity was subsequently estimated from ungulate carrying capacity and the effect of CC on the carnivoran faunas was analyzed in 12 paleocommunities from Southern Europe. Our results show that carnivoran species richness is strongly related to ungulate carrying capacity in recent ecosystems, but the late Early Pleistocene paleocommunities from Southern Europe included much richer carnivore guilds than would be expected for a recent community with a similar ungulate carrying capacity. Thus, those late Early Pleistocene ecosystems supported a high number of carnivoran species, but the carnivoran biomass they could support was relatively low. Consequently, carnivorans occurred at low densities in Southern Europe compared to the recent African savanna ecosystems, but likely also compared to coeval East African ecosystems. Consequently, the first Homo populations that arrived in Europe at the end of the late Early Pleistocene found mammal communities consisting of a low number of prey species, which accounted for a moderate herbivore biomass, as well as a diverse but not very abundant carnivore guild. This relatively low carnivoran density implies that the hominin-carnivore encounter rate was lower in the European ecosystems than in the coeval East African environments, suggesting that an opportunistic omnivorous hominin would have benefited from a reduced interference from the carnivore guild.
Chapter
The hyaena Chasmaporthetes lunensis lacks the robust and hyper-robust dentition of its bone-cracking relatives. Its gracile skeleton reflects an adaption to fast running and chasing down prey. The species was widespread on the northern hemisphere throughout the Pliocene and the Early Pleistocene ca. 4.5–1.5 myr, and, hence, an important component of the Holarctic fauna as a cursorial carnivore. By the earliest Pleistocene the relatively long-lived and seemingly fairly stable carnivore guild of the European Pliocene turned over, and canids of genera Lycaon and Canis replaced Chasmaporthetes. Its last known European occurrence is that from the German site Schernfeld with five isolated teeth documenting one large individual.
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The aardwolf Proteles cristatus is the only known hyaenid, living or extinct, to exhibit an extremely reduced dentition related to its termite-specializing diet. The fossil record of extant aardwolves extends to 2 to 4 million years ago, but records that inform its evolutionary origins are essentially nonexistent. Such circumstance renders it difficult to place this unusual hyena in the broader evolutionary context of small-bodied hyaenid species in Eurasian Neogene deposits. Here we describe a new genus and species of a small-bodied hyaenid, Gansuyaena megalotis, representing the closest morphological link to aardwolves to date. This new fossil hyena is based on a skull with associated mandible, a rostrum preserving several teeth, and several referred specimens. The new specimens were discovered in Neogene deposits in Linxia Basin, Gansu Province, China. Phylogenetic analysis indicates that among early hyaenids, G. megalotis is most closely related, but unlikely ancestral, to the living aardwolf. Also recognized in this new species are the fossils previously referred to "Protictitherium" aff. P. gaillardi from Pasalar, Turkey. Additionally, "Plioviverrops" guerini from Los Mansuetos, Spain is interpreted to represent a second Gansuyaena species. In addition to the living aardwolf, Proteles cristatus, our analyses suggest that the proteline lineage includes the extinct genera Gansuyaena, Mesoviverrops, and Plioviverrops. Although the precise timing and geographic location of evolutionary divergence between the aardwolf and Gansuyaena remain elusive, critical new morphological information provided by Gansuyaena specimens reinforce findings from recent genomic analyses that the aardwolf lineage has an ancient origin from small-bodied stem hyaenids prior to the appearance of large and robust bone-cracking hyaenines.
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A complete and well preserved fossil skeleton of Adcrocuta eximia (Roth & Wagner) is described. The skeleton consists of 156 bones. The locality is near the town of Hadzhidimovo, Blagoevgrad district, dated back as Late Maeotion, Turolian faunistic unit, MN12 zone. Comparisons are made with the skeleton of Chasmaporthetes borissiaki (Khomenko, 1931). It is concluded that it was an immature individual whose characteristics strongly correspond to Adcrocuta eximia. The differences found do not contradict to the taxonomical assignment.
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Hyenas (family Hyaenidae), as the sister group to cats (family Felidae), represent a deeply diverging branch within the cat-like carnivores (Feliformia). With an estimated population size of less than 10,000 individuals worldwide, the brown hyena (Parahyaena brunnea) represents the rarest of the four extant hyena species and has been listed as Near Threatened by the IUCN. Here we report a high coverage genome from a captive bred brown hyena and both mitochondrial and low-coverage nuclear genomes of 14 wild-caught brown hyena individuals from across southern Africa. We find that brown hyena harbour extremely low genetic diversity on both the mitochondrial and nuclear level, most likely resulting from a continuous and ongoing decline in effective population size that started about one million years ago and dramatically accelerated towards the end of the Pleistocene. Despite the strikingly low genetic diversity, we find no evidence of inbreeding within the captive bred individual and reveal phylogeographic structure, suggesting the existence of several potential sub-populations within the species.
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The majority of spotted hyena studies are conducted in places such as national parks and reserves where there are few humans present other than the researchers. I argue that this reflects a perception that “real” hyenas are those largely unaffected by contact with humans. This is at odds with fossil evidence which demonstrates a long, shared history of human/hyena co-evolution since our ancestors first came together on the African continent more than six million years ago. From that time human ancestors adopted lifeways that brought them into direct competition with hyena ancestors over carcass-based resources. These relations of competition and coexistence persisted through dispersals across Eurasia and into the late Pleistocene. So too in Africa, our respective ancestors competed over prey for millions of years. There hyena/human competition over livestock animals is a vestige of ancient enmity that marks both species as enemies. In light of this evidence I present a reconfiguration of what it is to be hyena or human. Using a theoretical framework developed by Jacob von Uexküll I argue that hyenas are to a great degree human-like and vice versa. This conclusion in turn undermines human exceptionalism by undermining the “non-animalness” which is normally held to separate humans from other species.
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19 taxa of carnivores are described from the Turolian (MN 11, Late Miocene) of Dorn-Dürkheim 1 (SW-Germany). These are Simocyon sp. (Mustelida inc. sed.), the meline Taxodon sp., the mustelines Eomellivora wimani, Promeles palaeatticus, Baranogale cf. adroveri, ?Circamustela sp., Martes cf. sansaniensis, one more uncertain species of Martes, and two musteline tooth fragments. Further the hyaenids Adcrociita eximia, Protictitherium crassum, Thalassictis robusta, the percrocutids Allohyaena kadici and Dinocrocuta sp., and the felids Felis attica, Paramachaerodits orientalis, Paramachaerodns ogygins, and Machairodus cf. aphanistus are found. The paleobiogeography and the phylogeny of the mentioned carnivores is discussed.
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The Agenian mammalian fauna (MN 1 and MN 2) has long been considered as a reduced Oligocene fauna because several mammalian groups as the theridomorph rodents, the creodonts of the genus Hyaenodon or the anthracotherids Anthracotherium and Microbunodon disappear at the level MN 1. A study of small carnivorans unearthed from Laugnac (Lot-et-Garonne, France) reveals the presence of both genera Semigenetta and Plioviverrops. S. laugnacensis belongs to a lineage whose size increases regularly during the Miocene with S. elegans, S. sansaniensis and S. steinheimensis. There is a discontinuity in the early Upper Miocene with the species S. grandis (punctuation of migration?). Another lineage composed with smaller species evolves also during the Miocene (S. cadeoti and S. ripolli). These genera are the forerunners of the Miocene migrants into the Aegenian age of mammals. The study of these genera shows also that the splitting between the extant carnivoran Feliformia families (Felidae, Viverridae, Hyaenidae and Herpestidae) must be dated from at least the Oligocene. -from English summary
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In the fossil record hyaenids are both diverse and abundant, and nearly 100 species have been named. Hyaenids have an expanded ectotympanic and semi-recumbent to recumbent septum bullae, and all fossil taxa likely to display this morphology are reviewed herein. The systematics and cladistics of the group are described. The age of the Hyaenidae is suggested to be some 25 million years. This is compatible with biochemical dates. Macroevolutionary patterns in the Hyaenidae are examined on the basis of the established cladogram and are found to be overwhelmingly gradual. Taxic patterns show that the Hyaenidae were most diverse in the late Miocene. -from Authors
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Evolution in the guild of larger carnivores is a marked feature of the Villafranchian faunal span. The eventual extinction of archaic forms and the incursion of taxa of more modern aspect finally produced a Middle Pleistocene guild similar in actual species, and almost identical in structure, to that of modern-day eastern Africa. However, the extinctions and appearances of members of the guild cannot be seen as a single tum-over, since the timing of the various events is both chronologically and geographically more complex. At around 1.6 Ma the hyaena Chasmaporthetes Iunensis became locally extinct and was replaced by Pachycrocuta brevirosfris, Panthera gombaszoegensis and several species of Canis. The large Pliocrocuta perrieri may also have become extinct around this period, although the timing is not clear. The dirk-toothed Megantereon cultridens managed to continue until the latest Lower Pleistocene at Untermassfeld, a time when Panthera leo, Panthera pardus and perhaps Crocuta crocuta dispersed into Europe. This incursion was followed during the earliest Middle Pleistocene by the re-appearance of Pliocrocuta perrieri. Towards the end of the Villafranchian, and until well into the Middle Pleistocene, the guild reached its greatest size and structural complexity with archaic taxa such as Homotherium latidens, Acinonyx pardinensis, Panthera gombaszoegensis, Pachycrocuta brevirostris and Canis (Xenocyon) lycaonoides coexisting with modern species such as Panthera leo, Panthera pardus, Crocuta crocuta and the small Canis mosbachensis. Only after 0. 5 Ma did the guild reduce in size with the disappearance of the archaic species. Geographic variability is marked by an early disappearance of Pachycrocuta brevirostris from Iberia and by the apparent absence of Acinonyx pardinensis after its last appearance at Puebla de Valverde. The latter species is now known to have reached Britain, based on a dental specimen from Norwich Crag Formation deposits at Easton Bavents. The reappearance of Pliocrocuta perrieri during the Middle Pleistocene did not apparently include Italy, Iberia or Britain, although both of the latter regions may have witnessed the latest appearances of Panthera gombaszoegensis. However, it is clear that many details of the biogeographic pattern of the guild remain to be established.
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In this beautifully illustrated natural history that links extinct larger feline species with those still in existence, collaborators Alan Turner and Mauricio Anton weave together the evidence of modern feline behavior with that of the fossil record. Turner's clear, insightful prose and Anton's masterly illustrations combine to offer specialists and newcomers alike an accurate and accessible guide to the evolution of cats.
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The holotype of the lower-middle Pliocene hyaenid Lycyaenops rhomboideae is redescribed and compared with contemporaneous hyaenids of the genera Lycyaena, Hyaenictis, Chasmaportheles and Pliocrocula. These comparisons show that the material represents a valid and distinct genus and species. The genus Lycyaenops is referred to the Chasmaporthetes lineage on the basis of its two-cusped ml talonid with reduced entoconid, reduced posterolingual cingulum cusp on p4 and premolar accessory cusps set in a straight line. It is distinguished from all other genera of that lineage by its smaller premolar accessory cusps, broad premolars and squared-off and very broad posterior premolar shelves. The species L. silberbergi, previously assigned to Chasmaporthetes, is also referred to Lycyaenops. It differs from L. rhomboideae in its greater development of the premolar accessory cusps and less developed posterior premolar shelves, but shares the broad, squared-off premolars. The interrelationships of Hyaenictis, Chasmaporthetes and Lycyaenops are at present best described by an unresolved trichotomy, with Lyeyaena as its sister taxon.
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The guild of African larger carnivores (Felidae, Hyaenidae and Canidae) that existed between c. 4.0 and 1.0 Ma had twice the number of species that exist there today. Lion and leopard appear to have originated in Africa soon after 4.0 Ma, and were confined to that continent until they appeared in Galerian faunas of Europe around 0.9 Ma. Cheetahs or cheetah-like forms were widely dispersed in both the Old and New World from the mid-Pliocene onwards, and their origins remain unclear. Spotted hyaenas are first known in Africa just after 4.0 Ma and appear in Europe in Galerian faunas, although they may have made an earlier dispersion into Asia. Brown and striped hyaenas appear to have originated in Africa by about 3.0 Ma. The former may be conspecific with the Eurasian Pachycrocuta perrieri, but while the latter appears to have dispersed into Asia during the Pleistocene, records of its presence in Europe require revision. Three other hyaenid species, P. brevirostris, Chasmaporthetes nitidula and C. silberbergi, along with archaic cats of the genera Homotherium, Megantereon and Dinofelis, are known from African Mid Pliocene-Lower Pleistocene deposits and also appear to have been part of an Afro-Eurasian dispersion. The pattern of evolution in the carnivore guild has broad correlations with the patterns of climatic change and the evolution of potential prey species.
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Fossil finds of the hyaenid genus Chasmaporthetes are reviewed. We consider the characters distinguishing this genus from Euryboas invalid and synonymize the two genera. The following species are included within the genus Chasmaporthetes: C. borissiaki (Khomenko), C. lunensis (Del Campana), with two subspecies, C. l. lunensis, from Europe, and C. l. honanensis, from China, C. nitidula (Ewer), C. ossifragus Hay, with two subspecies, C. o. ossifragus from the western United States, and C. ossifragus, an unnamed new subspecies from Florida, and C. exitelus, a new species from the Turolian of China. The presence of sexual dimorphism in C. lunensis is indicated from canine size. The stratigraphie range of the genus is early Blancan-Irvingtonian (North America), Turolian-Villa-franchian (Eurasia), Langebaanian to lower Pleistocene (Africa). The possible relationships of Chasmaporthetes are considered, and it is concluded that the genus is most closely related to the genus Thalassictis.
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The generalized ecological tolerances of larger terrestrial carnivores, reflected in their wide distributions in fossil and living faunas, make them good potential indicators of the changing pattern of links between Britain and the continent of Europe over time. Extensive contact is indicated until the earlier part of the Middle Pleistocene, that is prior to the Anglian-Elsterian glaciation. The pattern after that is more complex, with evidence for some form of marine barrier by the time of the Hoxnian and for complete isolation during the Last Interglacial.
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The large «crocutoïd hyaenas from the Plio-Pleistocenedeposits of Eurasia do not belong to the same lineage as the extant species Hyaena hyaena and must be referred to the genus Pachycrocuta Kretzoi, of which an emended diagnosis is given. This revision takes into account some undescribed or ill-known specimens from Russia, China and Africa. They definitely establish that Pachycrocuta ranged over Eurasia, from West Europe to East China, as early as the early Villafranchian at least, and that it was present in North Africa too; a large sample from the Odessa Catacombs affords an estimation of the intraspecific variation in the Ruscinian species, H. pyrenaica, known until now by only a few specimens from the western part of the Mediterranean basin, and it shows H. pyrenaica to be the ancestral form of the Villafranchian Eurasian species P. perrieri, from which derived P. brevirostris, the last species of the lineage, as previously shown by other authors. It appears that the "Hyaena lineage evolved simultaneously in Africa; we knew already that the root of this lineage is H. abronia, a species from the late Miocene of South Africa whose generic attribution is discussed relative to some Ictitheres from Shan-Si, Samos, Sahabi and Klein Zee. The hypothesis of a common African origin of the two lineages is not excluded, if not demonstrated. The relationships of the Pleistocene European species H. prisca and that of the extant African species H. brunnea are discussed.
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The Carnivores (Mammalia) from the Lower and Middle Pleistocene sites of Atapuerca (Spain). The Sierra de Atapuerca contains several karstic deposits rich in carnivores: Trinchera Galería, Trinchera Dolina, Trinchera Elefante and Sima de los Huesos. These remains do not belong to the same chronological period but to different moments of the Early and Middle Pleistocene. This complementary information provides a wider knowledge of the biostratigraphical carnivore sequence in Europe. Sima de los Huesos and Trichera Galería, although with different causes of accumulation which display differences on taphonomical analyses, contain similar species of carnivores which are typical Middle Pleistocene post-Cromerian taxa. This same period seems to correspond to the upper levels of Trinchera Dolina (TD11, TD10 and TD8b) anyway as the excavation works have not concluded yet, the fossil material recovered up to now is somewhat scarce to establish a definite conclusion about the chronology and type of carnivore community of these levels. The lowermost levels of Trinchera Dolina (TD3/4, TD5, and TD8a) contain a different carnivore assemblage with typical late Early Pleistocene-Cromerian species. This radical substitution of taxa is placed in the TD8 layer probably due to a stratigraphical hiatus in this level or in TD7. Trinchera Elefante is in a preliminary phase of study: the carnivores taxa recovered as yet differ than those from the lowermost levels of Trinchera Dolina suggesting this to be the most primitive association of carnivores in the Sierra.
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This study deals with the «percrocutoid hyaenids,present in Eurasia and in Africa in faunas of late Middle Miocene (Astaracian) to terminal Miocene (Vallesian, Turolian) age. These hyaenid species are characterized by certain dental apomorphies (loss of M2/2, specialization of the carnassials P⁴/M1, tendency for the hypertrophy of the anterior premolars) which differentiate them from other hyaenids with which they are found associated. In recent literature, these hyaenids have been attributed to the genera Percrocuta Kretzoi and Adcrocuta Kretzoi, the latter of which is monospecific. In fact some should be removed from the genus Percrocuta and assigned to the genus Allohyaena Kretzoi, itself subdivided into the two subgenera, Allohyaena Kretzoi and Dinocrocuta Schmidt-Kittler. The genus Adcrocuta is retained and the characters which distinguish this genus from Hyaenictis Gaudry are discussed.
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1996. The giant hyaena Pachycrocuta brevirostris (Mammalia, Carnivora, Hyaenidae). [L2ay~ne gSante, Pachycrocuta brevirostris (Mammalia, Carnivora, Hyaenidae)]. GEOBIOS, 29, 4 : 455-468. Villeurbanne le 31.10.1996 Manuscrit ddpos~ le 28.03.1995 ; accept~ d~finitivement le 29.05.1995. ABSTRACT -The giant hyaena, Pachycrocuta brevirostris, is known from deposits in Africa and Eurasia between ca. 3.0 and 0.5 Ma. It is the largest of the true hyaenas, but that size is not reflected in overall shoulder height since the distal limb segments are relatively shorter than those of living taxa. Its bodily proportions therefore appear to be suited to power and strength rather than speed. Its eventual extinction is part of the overall evolution in large carnivore guilds throughout the world and may have been closely linked to the extinction of machairodont cats. KEYWORDS : CARNIVORA, PACHYCROCUTA BREVIROSTRIS, PLIO-PLEISTOCENE, PALAEOECOLOGY. RI~SUMt~ -L'hy~ne g~ante, Pachycrocuta brevirostris, est connue des gisements africains et eurasiens entre 3,0 et 0,5 Ma. C'est la plus grande des hy~nes v~ritables, mais cette taille n'est pas refl~t~e dans la hauteur de l'~paule parce que les ~l~ments distaux des membres sont relativement courts en comparaison des hy~nes actuelles. Les proportions du corps sugg~rent un animal puissant mais rapide. L'extinction fait partie de l'~volution globale des groupes (guildes) de grands carnivores, et peut avoir ~t6 li~e h rextinction des f61id~s machairodontes.
Article
The topotypic material of the giant Late Miocene hyaenid Allohyaena kadici Kretzoi is described. New data on the deciduous dentition shows unambiguously that A. kadici is a hyaenid and not a percrocutid as reported by some previous authors. A. kadici is compared to the large hyaenids Adcrocuta eximia and Crocuta crocuta. These comparisons show that A. kadici has a mixture of primitive characters such as dp4 morphology, retention of m2, long and slender premolars and a large protocone on P4, and derived characters such as a preparastyle on P4, an internal root on P3 and a uniquely derived talonid structure of ml. This combination of features makes A. kadici difficult to classify, but it is considered to probably be most closely related to derived, bone-cracking hyaenids such as Pachycrocuta and Crocuta. A. kadici is rare in the fossil record, being found at only two sites. We suggest that the reason for this rarity is that it had a geographic and stratigraphic range which is poorly sampled in the Miocene fossil record of Europe.
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Material of the Miocene hyaenid Adcrocuta eximia from China is analysed statistically. No heterogeneities were found within this material. Comparisons with material from Samos and Pikermi, Greece, show that no taxonomic differentiation between these three samples is warranted. Adcrocuta eximia latro from the Sivalik deposits is provisionally considered a valid subspecies. The species A. australis from Langebaanweg, South Africa is removed from Adcrocuta to the genus Chasmaporthetes. The phylogenetic position of Adcrocuta has been subject to dispute, and for this reason we present a study of the interrelationships of selected hyaenid taxa using numerical cladistic methods. Two equally parsimonious trees of length 47 and consistency index 0.766 were found. Adcrocuta is placed as a sister-group to the Recent Crocuta crocuta, and not as a separate clade as suggested by other workers. Recent hyaenids form a crown-group which does not extend deep into the cladogram. Hyaena hyaena and H. brunnea are not sister-groups, and we resurrect the genus Parahyaena for the latter species.
Article
The holotype of the lower-middle Pliocene hyaenid Lycyaenops rhomboideae is redescribed and compared with contemporaneous hyaenids of the genera Lycyaena, Hyaenictis, Chasmaporthetes and Pliocmcuta. These comparisons show that the material represents a valid and distinct genus and species. The genus Lycyaenops is referred to the Chasmaporthetes lineage on the basis of its two-cusped m 1 talonid with reduced entoconid, reduced posterolingual cingulum cusp on p4 and premolar accessory cusps set in a straight line. It is distinguished from all other genera of that lineage by its smaller premolar accessory cusps, broad premolars and squared-off and very broad posterior premolar shelves. The species L. silberbergi, previously assigned to Chasmaporthetes, is also referred to Lycyaenops. It differs from L. rhomboideae in its greater development of the premolar accessory cusps and less developed posterior premolar shelves, but shares the broad, squared-off premolars. The interrelationships of Hyaenictis, Chasmaporthetes and Lycyaenops are at present best described by an unresolved trichotomy, with Lycyaena as its sister taxon.
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The Late Miocene (Late Vallesian, MN 10, about 9 Mya) carnivore trap of Batallones-1 (Madrid, Spain) has yielded a large sample of two species of sabre-toothed cats: the puma-sized Paramachairodus ogygia and the tiger-sized Machairodus aphanistus. This has allowed, for the first time, complete studies of the biomechanics and comparative anatomy of these animals. Focusing our study on the small species, Par. ogygia, the most richly represented and best known carnivore from Batallones-1, we attempt to infer some aspects of the behaviour and ecology of this early sabre-toothed cat, such as breeding behaviour, the degree of social interaction between individuals, sexual dimorphism, preferred habitat and prey size. Our results suggest that Par. ogygia was a solitary felid with a low sexual dimorphism index, which in turn indicates low competition between males for access to females, and some degree of tolerance between adults, so that young adults were allowed to share the territory of their mothers for some time after maturity. The machairodont adaptations of Par. ogygia indicate that this species was able to subdue and kill prey in less time than pantherines do, thus minimizing the risk of injury and the energetic costs of this action. In a wider context, the carnivore guild of Batallones-1 and the overall mammal community indicate that the landscape around the trap was a wooded habitat. Batallones-1 is thus establishing itself as one of