이 게시 글 상당히 길 고 . . 취향 에 맞 지 않 는 분 이 많이 계실 듯 하 니 그런 분 들 은 그냥 뒤 로 가 기 눌러 주 시 면 됩니다 . 소개 될 동물 들 의 목록 입니다 . 그리고 저걸 다 찍 기 엔 너무 많 아서 캡쳐 한 사진 파일 일부 와 그리고 더 많 은 내용 을 텍스트 로 옮겨 나열 하 겠 습니다 . 그리고 물론 순서 대로 나열 하 겠 습니다 . 먼저 개과 입니다 . 아 그리고 The dental formula of Carnivora is I 3 - i 3 ; C 1 – c 1 ; P 4 – p 4 ; M 3 - m 3 p 4 나 m 1 같 은 단어 가 보이 실 텐데 여시 서 말 하 다 시피 이빨 쪽 들 의 길이 를 나타내 는 것 입니다 . m 1 은 아래쪽 열 육 치 p 4 는 위쪽 열 육 치 . . P 3 는 소구 열 육 치 . . 저런 이빨 길 이 및 두 개 골 길이 들 등 으로 몸무게 를 추산 해 내 는 것 입니다 . 후 , , 영어 도 못하 고 내용 도 너무 많 아서 아주 간단 하 게 몸무게 같 은 정보 만 말 하 고 넘어가 겠 습니다 . Nyctereutes megamastoides ( Pomel , 1842 ) This racoon dog is poorly represented in the Italian Plio - Pleistocene and appears only in the PCOMMontopoli ( localities Montopoli and Colle Pardo ) . This species is probably replaced by the arrivalof wolf like Villafranchian canids all over Europe and it characterizes the fauna of middleVillafranchian . Thorough anatomical descriptions of the European forms of this species are in Viret ( 1954 ) : the skull is characterized by strong sagittal and nucal crests , the anterior part of the palate is narrow and slender and becomes elongated posteriorly ; the mandible presents a typical subangular lobe ( Fig . 1 . 3 ) ; dentition is fox - like with short canines and an elongated carnassials even if sometraits are discriminant ( e . g : size of M 2 relative to P 4 ) and clearly separate the genus Nyctereutesfrom Vulpes ( Koufos 1993 ) . A recent description of long bones morphology is known from the material of Saint Vallier : generally it resembles the proportion of the genus Vulpes even if some bones are wider on thearticular surface ( e . g . the humerus and radius ) , the tibia presents a typical morphology in the crest ( Argant 2004 ) . Generally , the morphology of Villafranchian raccon dog resembles that of itsmodern counterpart . On the basis of this observation Kurtén ( 1968 ) suggested an adaptation toomnivory feeding habits dominated by vegetables and sometimes meat . As the Italian fossil material is fragmented I used for morphometric analysis several completemandible samples from IGME ( Istituto Geológico y Minero de España . Museo Geominero . Madrid , Spain . ) and NMB ( Naturhistorisches Museum Basel , Switzerland ) . The estimated body mass shouldbe usually larger than that of modern racoon dogs ( Nyctereutes procyonoides ) ( Kurtén 1968 ) . Onthe basis of several lower carnassial measurements is 7 . 7 kilograms but a complete skull from Saint Vallier ( Argant 2004 ) gave a better estimated body mass of 11 kilograms that will be considered valid and more accurate for this thesis . Nyctereutes megamastoides , 즉 너구리 의 옛 조상 의 몸무게 는 carnassial 길이 들 을 측정 결과 7 . 7 킬로 로 나왔 다 하 며 11 킬로 에 까지 이를 수 있 다 말 합니다 . The post cranial skeleton morphology is similar to that of C . lupus with size proportion resembling small wolves . Generally , Canis etruscus is like a shepherd dog in size ( Kurtén 1968 ) while p a l a e o ecologicalreconstructions have been performed on the basis of Venta Micena ( Spain ) specimens ( Palmqvist etal . 1999 ; 2002 ) . The Spanish fossil material belong to a much smaller form than Italian and Rook ( 1993 ) proposed that it belong to Canis aff . arnensis . For this reason , here , I use a body massreconstructed only from Italian fossil specimens . According with Meloro et al . ( 2007 , Appendixdata ) the estimated body mass of C . etruscus is 21 kilograms . Palmqvist et al . ( 1999 , 2003 ) suggested an omnivorous diet but there are no evidence from Italian specimens . Usually Canis etruscus is considered a pack hunter but there is no formal evidence or palaeoecological reconstruction of such a behaviour . It is worth mentioning that C . etruscus is one of the most abundant canids in Italy ( Raia et al . 2006 b ) and , probably , Europe as well . Palaeoecology of the Etruscan wolf will be clarified in several morpho - ecological analyses based on the mandible and long bones measurements ( see Chapters 3 – 5 ) . The stratigraphic range of this species is restricted to Upper Villafranchian . In Italy , C . etruscus is present in successive Montopoli , Upper Valdarno and ValdiChiana PCOMs . Canis etruscus , 현재 늑대 의 조상 늑대 는 몸무게 가 약 21 킬로그램 정도 였 다 합니다 . Canis arnensis ( Del Campana , 1913 ) The taxonomic history of this species is complex and it is necessary to elucidate several points to clarify the position adopted in this thesis . The description of this species comes from the abundant material of Valdarno ( Del Campana 1913 ) and it was successively analysed by Torre ( 1967 ) and Rook ( 1993 ) . Rook ( 1993 ) proposed a new species similar in morphology to C . arnensis but more advanced in the dentition . This new species is interpreted as Canis aff . arnensis and its stratigraphic range is limited to the middle Pleistocene ( Galerian ) of the Mediterranean region ( Rook and Torre 1996 b ) . The authors argue for caution of this interpretation because a comparison with the material of Galerian dogs of Central - North Europe was not performed . Conservatively , here I considered all the uncertain nomenclature as Canis arnesis = Canis cfr . arnensis = Canis aff . arnensis in keeping with Raia et al . ( 2005 ; 2006 a ; 2006 b ) as belonging to C . arnensis . Generally , Canis arnensis resembles jackals in skull morphology with a short muzzle ( compared to C . etruscus ) , poorly developed nasals and frontal sinus and a weak sagittal crest ( Torre 1967 ) . The teeth ratio were interpreted as jackal - like but Kurtén ( 1974 ) points out a stronger affinity with coyotes in M 1 / P 4 , p 4 / m 1 and m 1 length / m 1 breadth ratios . Canis aff . arnensis has a broader upper M 1 and a shorter trigonid in m 1 compared with C . arnensis ( Rook 1993 ) , but the taxonomic relevance of such features is unclear . Post - cranial fossil material of Canis arnensis is scanty and fragmented but Torre ( 1967 ) stressed how the remains of Valdarno resemble modern jackals in proportion and morphology . The palaeoecology of this canid was never clarified , although Kurtén ( 1974 ) suggested a closeaffinity with the North American coyote ancestor Canis priscolatrans indicating that C . arnensiscould be a possible geographic variant of a large coyote holartic population . Body mass estimates is reported in Meloro et al . ( 2007 ) as 16 kilograms and will be used for this thesis . Its stratigraphic range is the largest ( especially in virtue of the uncertain taxonomic validity of Galerian forms ) . Its presence in PCOMs units span from Upper Valdarno ( Upper Villafranchian ) to Aurelian with some advanced forms recorded at Capri ( Quisisana ) , Contrada Monticelli , Grotta Romanelli ( lower beds of “ terre rosse ” ) and Campo Verde ( Raia et al . 2006 a ) . Canis arnensis , 코요테 와 비슷 하 다는 이 동물 의 체중 은 나오 질 않 네요 . From a palaeoecological perspective , L . falconeri is well known ( Palmqvist et al . 1999 ; 2003 ) . Acomplete skull from Venta Micena has anomalies that could have severely affected the survival of the individual it belongs to . This fact , together with further anatomical analyses , suggests abehavioural and morpho - ecological similarity between L . falconeri with the modern African wilddog . Probably this highly - cursorial , pack hunting species was capable to kill horses ( e . g . Equus stenosis ) and ungulates adapted to open terrains ( Palmqvist et al . 2003 ) . The estimated body mass should be 28 kilograms ( Palmqvist et al . 1999 ) but Meloro et al . ( 2007 ) reported 26 kilograms on the basis of Italian fossil material in Rook ( 1994 ) . The latter value ( that is not so different from Palmqvist et al . 1999 ) will be considered valid . Lycaon falconeri is present in Upper Valdarno and Pirro Nord PCOMs . Its absence fromValdiChiana PCOM suggests rarity in this species for a short period of the Italian fossil record . Lycaon falconeri , 유럽 리카온 은 체중 이 약 28 킬로그램 에 달 한다는 것 과 다른 학자 는 화석 들 로 추산 해 낸 결과 가 26 킬로그램 이 라는군요 . Canis lupus , 회색 늑대 의 경우 지역 에 따라 다르 지만 평균 38 킬로 정도 . 음 . . 한마디 로 고대 나 지금 이나 체중 이 비슷 했 다는 이야기 인가 . . 그리고 다음 은 mustelidae , 족제비 과 입니다 . Meles meles ( Linnaeus , 1758 ) This form has a unique omnivore morphology and it has been recorded in few but significant sites of the Italian Paleo - Communities : in the locality G . R . A of the Galerian 2 ( Caloi and Palombo 1986 ) Visogliano of Galerian 3 and Torre in Pietra of the Aurelian ( Data source Palombo et al . 2003 ) . In Slivia of Galerian 1 Palombo et al . ( 2003 ) report cf . Meles meles but this record , which should be the oldest for the Italian peninsula , has not been considered because it is doubtful . Even if it is rare , the badger is present continuously from the Galerian to Aurelian . It is a typicalmember of modern Holarctic fauna and Kurtén ( 1968 ) reported first occurrences of the species allover European localities from the middle Pleistocene . The skull is massive with a strong sagittal crest , the rostrum is short relative to the braincase as it is common in mustelids ; the lower carnassial has an elongated talonid region relative to the trigonid and with some accessory cusps . The post cranial morphology is typical as in burrowing animals with short legs and large carpals and tarsals . The considered Italian fossil material is represented by few fragments and for morphometricanalyses I used fossil material of the Italian Late Pleistocene from Monte Rufeno ( Viterbo , Italy ) preserved in the Pigorini Museum ( Rome ) . In the described material from G . R . A . there are nosignificant differences in size and morphology from extant badgers ( Caloi and Palombo 1986 ) and for this reason a body mass of 12 kilograms ( cf . Gittleman 1985 ) was assumed . Meles meles , 유럽 오소리 의 경우 추정 해 낸 체중 이 12 킬로 라 합니다 . 다음 은 곰과 입니다 . Ursus minimus ( Devèze and Bouillet , 1827 ) The most important Italian remains of Ursus minimus came from Gaville Valdarno and werereviewed by Berzi ( 1966 ) . She pointed out close similarities of U . minimus with modern black bear ( U . thibethanus ) and such opinion has been validated in Mazza and Rustioni ( 1994 ) who grouped U . minimus and U . thibetanus . In the skull , the neurocranium appears more elongated than the rostrum , the upper premolars 1 - 3 are small and spaced like the lower premolars 1 - 3 , the lower carnassial exhibits a zig - zag morphology defined by the lingual and buccal cusps similar to that of modern black bears . The dentition does not show a strict trend in the p 4 size reduction because in U . minimus this tooth is smaller than in U . etruscus ( Ficcarelli 1979 a ; Fig . 1 . 6 a , b ) but it has beenclarified that U . minimus represents a primitive stock for the evolution of both European and Asiatic bears ( Mazza and Rustioni 1994 ; Petronio et al . 2003 ) . In the post - cranial elements Berzi ( 1966 ) reconstructed the correct physiological position of metapodials and metatarsals which should have allowed the species to climb the trees . Among Italian PCOMs , U . minimus is recorded only at Triversa with a very limited fossil record ( only two local occurrences ) . Palaeoecology of this form was never clarified and a body mass of 104 kilograms has been reconstructed by using skull - length ( in Meloro et al . 2007 ) . Mazza and Rustioni ( 1992 ) furnished a comprehensive overview of Ursus etruscus morphology as well as occurrence in Plio - Pleistocene Italian sites . This bear is restricted to the UpperVillafranchian and it occurs continuosly in PCOMs Upper Valdarno , Val di Chiana and Pirro . The morphology is peculiar and it confirms that premolar reduction in bears does not begin with the cave bears but earlier . The skull is elongated and flatted , premolars ( upper and lower ) are reduced ( Fig . 1 . 6 b ) , the lower carnassial and the upper M 1 exhibits a lateral compression . Such features suggests an early adaptation to omnivory but a thorough palaeoecological reconstruction has never been performed . Long bones are generally more elongated than in U . minimus with plantigrady adaptations such as in brown bear . A body mass reconstruction based on the Italian fossil specimens gave a value of 160 kilograms ( in Meloro et al . 2007 ) . Ursus minimus , 반달곰 의 조상 이 라는 이 곰 은 이탈리아 의 화석 들 로 추정 해 낸 체중 이 160 킬로 라는군요 . 호오 예전 유럽 엔 반달곰 이 . . 학명 Ursus deningeri , Deninger bear 라 불리 는 이 곰 은 동굴 곰 과 체중 이 비슷 했 다는 데 275 킬로 정도 로 추산 된다 하 며 Ursus spelaeus , 동굴 곰 은 두개골 로 추정 해 낸 체중 이 약 300 킬로 이상 에 달할 수 있 다 말 하 며 이 자료 저자 는 275 킬로 쯤 될 것 같 다 . . 라고 하 는군요 . 그리고 그 외 동굴 곰 의 뼈 로 추산 해 낸 체중 자료 가 또 있 는데 LIMB BONE PROPORTIONS AND BODY MASS OF THE CAVE BEAR ( URSUS SPELAEUS ) 입니다 . 거기 선 mass of the male cave bears of these Late Pleistocene populations was 319 kg , 라고 하 며 다른 자료 인 Sexual dimorphism and ontogenetic variation in the skull of the cave bear ( Ursus spelaeus Rosenmüller ) of the European Upper Pleistocene 에선 from 319 kg on average for males and 244 kg on average for females ( Viranta 수컷 평균 319 킬로 , 암컷 평균 244 킬로 라고 하 는군요 . Ursus arctos ( Linnaeus , 1758 ) The brown bear characterizes several assemblages of late Pleistocene in Italy . Its commonness in the Aurelian is low ( with just two occurrences ) and the earliest certain record is from Spessa 2 ( PCOM Galerian 3 ) . This species exhibits a large body size and , actually , it is spread all overcontinental Europe and North America with several sub - species . In Italy , brown bear persists with a small population in the central Appennine but it is recolonizing the Alps thanks to severalreintroduction programs . The most significant Italian remains were described by Capasso Babato et al . ( 1982 ) belonging to the Upper Pleistocene site of Monte Cucco ( not included in Aurelian fossil localities ) . Generally , the Pleistocene brown bear becomes larger in size than the actual forms but their morphology does not change so much . Its dentition is entirely adapted to an omnivorous feeding habit and this fact allows to consume meat as well ( with no extreme reduction as in U . deningeri - spelaeus group ) . The skull is less prominent than in cave bears and long bone morphology is generally more slender . As this species is extant , there are few palaeoecological reconstructions about some peculiar behaviours or more accentuated meat consumption in the diet of Pleistocene bears . Stiner et al . ( 1998 ) reported a high degree of overlap in the diet of U . deningeri and brown bear . Actually , the brown bear occupies forest habitat but also mountain high plains with shrub vegetation . Its diet is based on vegetables and insects but meat is consumed to some extent . The body mass range is really high and it depends on the regions : here I use an estimate of 167 kilograms based on the maximum skull length of specimens figured in Capasso Barbato et al . ( 1993 ) in Meloro et al . ( 2007 ) . 그리고 Ursus arctos , 불곰 의 경우 엔 167 킬로 로 추산 했 군요 . 다음 은 하이에나 류 들 입니다 . Chasmaporthetes lunensis ( Del Campana , 1914 ) The running hyena is a typical Villafranchian form and its earliest occurrence in Italy is at Triversa . Del Campana ( 1914 ) described the morphology of the palate but he identified the fossil material as belonging to the genus Lycyaena . Later , Viret and De Beaumont ( in Werdelin and Solounias 1991 ) re - interpreted this species as Euryboas lunensis . Kurtén and Werdelin ( 1988 ) reviewed the genus Chasmaporthetes clarifying the relevance of several dental features which allow to distinguish Euryboas spp . from Chasmaporthetes lunensis . The latter is characterised by the absence of p 1 and m 2 in the lower jaw which is slender and elongate ( Fig . 1 . 8 a ) ; the skull is Hyaena - like with a strong sagittal crest a broad rostrum but also a well - developed braincase ; the upper premolars are slender and elongate with the P 4 carnassial possessing a prominent protocone and a long parastyle ( Fig . 1 . 8 b ) . In the m 1 proto and paraconid are subequal in size and the talonid exhibits a well developed hypoconid with a blade function . Post cranial material is well represented only in the North American form C . ossifragus ( Berta 1981 ) which is characterised by typical cursorial limb proportion . On the basis of material from Mt . Perrier Kurtén and Werdelin ( 1988 ) observed that the European C . lunensis possesses a radius longer than the tibia . There is a complete metacarpal from Layna in Spain while the other long bones are poorly known . On the basis of some aspects of functional morphology Kurtén and Werdelin ( 1988 ) propose that C . lunensis was an active predator with some adaptations to bone cracking as well . Ferretti ( 1999 , 2007 ) confirms the presence of a typical zig zag structures in tooth enamel but in a more primitive ( as compared to other hyenas ) condition and , consequently , minor bone cracking specialization . The ecological rule of C . lunensis was probably similar to that of modern spotted hyena in the African savannah . The body weight of this animal ( 75 kilograms in Meloro et al . 2007 ) partially resembles that of a very large modern spotted hyena ( Kruuk 1972 ) . Pliocrocuta perrieri ( Croizet and Jobert , 1828 ) This species spread its range two times all over Europe : during the middle Villafranchian and in the Galerian ( Turner 1990 ) . In Italy it occurs in two localities of PCOM Montopoli , than it disappeared in Upper Valdarno until a single occurrence of a fragmented upper carnassial in Rome during the Galerian 2 ( Palombo et al . 2003 ; Caloi and Palombo 1986 ) . Morphological features have been well described by Howell and Petter ( 1980 ) : the skull has a relatively long muzzle , a short braincase with a well developed sagittal crest , the upper carnassial has a strong protocone while in the lower premolars there is an increasing size of premolars . The carnassial m 1 is diagnostic : it possesses a thick trigonid and a little - developed hypo - entoconid ; the metaconid is not present or poorly developed . This latter feature allows to distinguish P . perrieri with the modern brown hyena ( Parahyaena brunnea ) with which there is a remarkable morphological similarity ( Kurtén 1968 ) . Post - cranial material is unknown except for humerus fragments from Montopoli and St . Vallier that resembles modern Crocuta and Hyaena in its morphology ( Argant 2004 ) . The close similarity with modern brown hyena suggests a scavenging adaptation although there are no other evidences on behavioural preference to hunting or scavenging . Body mass reconstruction is based on m 1 length from Italian specimens ( Bebi 1998 ) and it gives a value of 80 kilograms ( Meloro et al . 2007 ) . Chasmaporthetes lunensis , running hyena 라 불리 는 녀석 인데 75 킬로 정도 의 몸무게 를 예상 한다 하 며 . . Pliocrocuta perrieri , Perrier 하이에나 불리 는 녀석 은 80 킬로 로 추산 한다 합니다 . Pachycrocuta brevirostris , 쇼트 페이 시드 하이 에 나라 불리 는 이 녀석 은 현재 암사 자만 한 덩치 인 127 킬로 의 체중 이 라 합니다 . 중간 에 Pliocrocuta perrieri 가 나와서 그림 끼워 봅니다 . 동굴 하이에나 보다 더 큰 덩치 를 보이 는 . . 다음 은 동굴 하이에나 입니다 . Crocuta crocuta , 뒤 에다가 spelaea 도 붙여야 하 는데 . . 곰 도 그렇 고 하이에나 , 사자 도 아 종명 에다가 spelaea 가 붙 으면 동굴 ~ 이라고 명명 하 면 되 더군요 . 어쨌든 동굴 하이에나 의 경우 엔 몸무게 를 102 킬로 로 추산 하 는군요 . 하이에나 과 다음 내용 인 . . 고양이 과 입니다 . but just the Layna ( Spain ) fossil samples . The evolutionary lineage of lynx is represented in theMiddle Pleistocene by the small Lynx pardinus like . The species name of such form has beenproposed according with its similarity to the extant Spanish endemic lynx , and Raia et al . ( 2006 b ) consider the only Italian record as L . pardina spelaeus . Recently , Testu ( 2006 ) argues that suchspecific name is not valid while the correct nomenclature to describe middle Pleistocene forms is Lynx spelaeus and this will be also applied here . During the middle Pleistocene the Villafranchian forms are replaced by modern pantherine catswhich spread all over Eurasia : Panthera leo and Panthera pardus . These forms will be much more common in Italian Upper Pleistocene deposits achieving high abundance ( e . g : in the Equi cave Del Campana 1947 ) . The Pleistocene leopard does not represent a distinct extinct species while for the lion several specific or sub - specific names have been proposed . Here , conservatively , I use the old Kurtén ( 1968 ) proposal about the fact that the Pleistocene lion were remarkably larger than extant one because of different climate but such feature is not enough to support a different species . The Italian lion from Isernia probably represent one of the oldest European specimen , even if , unfortunately , it is represented only by one tooth ( Sala 1990 ) . Acinonyx pardinensis ( Croizet and Jobert , 1828 ) The giant cheetah is well represented in the Villafranchian fossil record from Italy . This species occurs at Triversa , in two localities ( Montopoli and Costa S . Giacomo ) of Montopoli PCOM , at Olivola , Casa Frata and Pirro . Its geographic range covers all Europe since Asiatic localities as well ( Kurtén 1968 ) . Morphologically , Acinonyx pardinensis resembles its modern African counterpart A . jubatus and only size represents an evident trait of discrimination ( Ficcarelli 1984 ; Turner andAntón 1997 ; Argant 2004 ) . The skull possesses a distinct shape with high and enlarged nasals , high and well developed orbits , a broad rostrum in the posterior region . The upper dentition ischaracterised by short canines , a P 3 with a reduced anterior lateral cusp which is larger in A . jubatus and an upper carnassial with a more reduced paracone . The lower jaw of A . pardinensis is similar to that of A . jubatus and only the morphology of p 3 isdistinctive . Post cranial material is well known from the specimens ( hind limb bones ) collected at Olivola and several fore limb elements from St . Vallier . Their morphology is similar to that ofmodern cheetah and this fact suggests a similar adaptation ( probably more advanced ) to high - speed chasing . All the morphological traits lead considering the giant cheetah as similar in ecology to the modern forms ( Turner and Antón 1997 ) . The body weight estimates should resemble a lion - sized animal of c . ca 90 kilograms in keeping with Argant ( 2004 ) . For this thesis I will use a conservative estimate of 67 kilograms based on lower carnassial length ( Meloro et al . 2007 ) . Acinonyx pardinensis , 자이언트 치타 라 불리 는 녀석 이 지요 . 체중 은 90 킬로 에 도 달 할 수 있 다는 언급 과 이 자료 의 저자 는 67 킬로 정도 로 추산 하 고 있 습니다 . 뭐 그래도 여전히 현생 치타 에 비하 면 가장 크 다 할 수 있 습니다 . 다음 은 검치호 들 입니다 . ( see below ) . A peculiar pathology has been recorded on humeri from French localities ( St . Vallier and Senèze ) and probably this demonstrates how the fore limbs were overloaded during the chasing of prey ( Turner and Antón 1997 ; Argant 2004 ) . This cat was remarkable heavy and probably its body weight was greater then that of a large lion . Here , I use an estimate of 231 kilograms ( Meloro et al . 2007 ) . Homotherium latidens ( Owen , 1846 ) The late Homotherium is represented in three Galerian localities of the Italian fossil record ( Palombo et al . 2003 ; Raia et al . 2006 b ) . This species usually resembles H . crenatidens in skull morphology as well as post cranial elements even if the canines are shorter and differ in shape ( Ficcarelli 1979 b ) . The functional morphology of this species has been broadly investigated especially in the light of the long debate concerning its hunting behaviour ( Antón and Galobart 1999 ; Antón et al . 2004 , 2005 ) . Basically , there are two theories : the “ stabbing ” suggests that sabre tooth cats use their sabres for a lacerating function ( the attack to the prey should be at belly or other soft parts ) and this implies a rotation along the thoracico - cervical joint ; the “ canine shear bite ” implies a much more precision in the movement of the neck whose strong musculature allows toperform a precise jugular bite ( while the strong fore limbs stop the potential prey movements ) . The studies of Spanish palaeontologists agrees with this latter theory and widely demonstrate its validity on the basis of the well developed mastoid region together with cervical vertebrae morphology and lower jaw and dental features . Antón et al . ( 2005 ) demonstrated also that Homoterium latidens was broadly adapted to cursoriality and its fore limb is not so strong to allow a single individual in bringing down large ungulates . This fact suggests that H . latidens was constrained to hunt in group like modern lions do even if its hunting technique differs in immobilising prey as well as in the canine shearing biting . On the basis of Texas cave remains belonging to Homotherium serum , it has been suggested that also Homotherium latidens specialized its target prey on juvenile of mammoths ( Turner and Antón 1997 ) . Palmqvist et al . ( 1996 ) evidenced for H . latidens a prey class selectionsimilar to the extant lion . Later Palmqvist et al . ( 2003 ) also demonstrated that isotopic values ( carbon , nitrogen and oxygen ) of the sabre tooth cat at Venta Micena is similar to that of largegrazer ungulates and most interestingly to juvenile mammoths . There are no doubt that H . latidens was a cursorial predator and then specialised on grazer ungulates . The specialization on juvenile mammoths still to be debated especially on the basis of long bone morphology that does not support completely such a behaviour ( Antón et al . 2005 ) . The body mass of Homotherium latidens is higher than that of the Villafranchian H . crenatidens ( 274 kilograms ) . It is noteworthy that this reconstruction is based on lower carnassial length and it is only indicative but not accurate especially for felids whose body mass greatly variesintraspecifically . Homotherium crenatidens , 유럽 에 살 던 검치호 의 일종 으로써 체중 은 커다란 사자 정도 인 . . 231 킬로 로 추산 한다 합니다 . 그리고 Homotherium latidens , 역시 검치호 의 일종 으로써 앞 의 녀석 보다 더 큰 274 킬로 의 몸무게 로 보 고 있 군요 . . 이런 검치호 종류 들 은 오늘날 의 사자 처럼 무리지 어 사냥 했으며 큰 덩치 와 큰 송곳니 로 매머드 새끼 도 사냥 했 을 것 으로 예상 한다 합니다 . 다음 은 dirk - tooth cat . . 단치 호 ( ? ) 입니다 . Megantereon cultridens ( Cuvier , 1824 ) The dirk tooth cat is another characteristic member of the Italian Villafranchian . Turner ( 1987 ) furnished an accurate diagnosis of M . cultridens but he considered M . whitei in synonymy with the latter justified by the potential high sexual size dimorphism . Sardella ( 1998 ) reviewed the taxonomy of Megantereon spp . and grouped in M . cultridens also the Italian specimens from Pirro and Farneta . In Raia et al . ( 2006 b ) M . cultridens is present only in Montopoli and Upper Valdarno PCOMs but not Pirro . According with Palmqvist et al . ( 2007 ) I consider valid such strictstratigraphic range to M . cultridens present only in Valdarno FU ( = Upper Valdarno PCOM ) . Megantereon cultridens can be easily distinguished from Homotherium spp . for its smaller size and its longer non - crenulated canines . The canines are unserrated , more curved and laterally compressed . The brain case of M . cultridens is long while the rostrum is shorter and broad posteriorly . There is a short diastema between the incisors and the upper canine that in Homotherium is less developed . Both second premolars ( upper and lower ) are not present . The ratio between P 4 and P 3 is highly variable and in the P 4 the deuterocone is usually prominent . In the lower jaw , the canine is very reduced as well as p 3 . The presence of a developed mandibular flange is a distinctive trait ( Fig . 1 . 12 ) . The neck is long and the limbs are short and robust . Such features explains the peculiar hunting behaviour that should conform to canine shearing bite . In Megantereon the limb bones are stronger than in Homotherium and this feature allows thespecies to ambush its prey alone ( Turner and Antón 1997 ) . It is worth mentioning that post cranial material of M . cultridens is scanty ( Argant 2004 describes several metapodials ) and most of long bone morphology is better known for M . whitei . The size of M . cultridens is much smaller ( 63 kilograms in Meloro et al . 2007 ) than H . crenatidens but its morphology probably is indicative of a solitary hunting style . The target prey of Megantereon were represented by medium size ungulates but also large deer of the genus Eucladoceros ( Turner and Antón 1997 ) . Much morepalaeoecological reconstructions have been proposed for Meganteron whitei of Venta Micena . Megantereon whitei ( Broom , 1937 ) . This advanced form of Megantereon is recorded only at Pirro site in Italy ( Raia et al . 2006 b ) andother European localities Galerian in age like Untermassfield ( Germany ) and Venta Micena ( Spain ) ( Palmqvist et al . 2007 ) . Martínez - Navarro and Palmqvist ( 1996 ) underline the importance of this species in understanding the routes of Homo dispersal from Africa to Europe and Asia . Probably this hypercarnivorous species provide carcass for scavenging by early Homo members . Morphologically , M . whitei is similar to M . cultridens and can be distinguished on the basis ofseveral features : upper canine length and breadth ; P 4 , m 1 and p 4 length and breadth . The dirkcanines are much more developed than in M . cultridens while the carnassials are smaller in size . Interestingly , also the mandible shape differs among the two species with M . cultridens having adifferent biomechanical efficiency than M . whitei that maximized the bite force at the m 1 . Post cranial anatomy should resemble the extant jaguar ( Panthera onca ) according with the studies based on African Megantereon ( Lewis 1997 ) . Fore limbs are stronger and massive . Such trait are characteristic of an ambush predator and Palmqvist et al . ( 2003 ) confirms that M . whitei prefers habitat with cover vegetation and as consequence also browser ungulates . Although , the morphology of M . whitei is well equipped in killing prey much more larger that the predator itself . Limb morphology suggests also a slight adaptation in tree climbing but such behaviour isimplausible considering the long canines . Body weight reconstruction of this species differs a lot when using several traits . A body mass of 55 kilograms is estimated when using m 1 length or 100 kilograms when using humerus cross sectional diaphysis ( Martínez - Navarro and Palmqvist 1996 ) . Although the authors suggest a body mass of 100 kilograms is probably much more accurate consider valid a body weight of 55 kilograms that is much more similar to that of M . cultridens for which a reconstruction based on skull length ( PER 2001 a ) achieves a 60 kilograms value . Megantereon cultridens 단치 호 의 종류 인 이 녀석 은 체중 추산 이 약 63 킬로 이 며 . . Megantereon whitei 역시 단치 호 의 종류 인 이 녀석 은 100 킬로 에 도 달 할 수 있 다 하 며 평균 이 60 킬로 정도 로 추산 된다 하 는 거 같 군요 . Lynx issiodorensis ( Croizet and Jobert , 1862 ) The Etouaires lynx is another typical Villafranchian element that spread all over Europe . In Italy itis present continuously in Upper Valdarno , ValdiChiana and Pirro PCOMs ( Raia et al . 2006 b ) . The Valdarno fossil material was described in detail by Fabrini ( 1896 ) who furnished accuratedescription of both cranial and post cranial elements . But Kurtén ( 1978 ) furnished an accuratecomparative analysis of L . issiodorensis with modern linxes . The skull is very large with narrowzygomatic arches and wider and longer rostrum ( compared with the skull length ) ( Fig . 1 . 13 ) . The nucal crest is well developed and also lower jaw morphology suggests a strong biting force : mandible possess a large dentition and a deep masseteric fossa . On the upper dentition , the P 2 is not present , P 3 is slender and more elongated than in modern Lynx and in the upper carnassial a small ectoparastyle is prominent ( Fig . 1 . 13 ) . In the mandible there is a long diastema , p 3 and p 4 can be separated by a gap and m 1 is very short lacking the metaconid and talonid complex . L . issiodorensis exhibits a long neck as well as long lumbar region while the hind limb region is less developed than in modern lynx . Generally Kurtén ( 1978 ) noted that from the 4 th lumbar vertebra to the anterior part of the body the Etouaires lynx is larger than the extant lynx while from the 5 th lumbar vertebrae it is shorter . Limb bones proportion resembles the American puma ( Puma concolor ) being more robust than inlynx . In Europe Werdelin ( 1981 ) suggests an evolutionary trend in L . issiodorensis from earlyVillafranchian members that are larger in size to the late representatives of the species that aresmaller but possess a longer m 1 . Palaeoecological reconstruction of L . issiodorensis were deduced from its morphological similarity with modern lynx : probably the Etouaires lynx specialized on small rabbits ( e . g : Oryctolagus spp . ) but was also capable of killing medium size ungulates ( like the modern European lynx kills roe deer ) ( Kurtén 1978 ) . A body mass reconstruction of 22 kilograms is here applied on the basis of m 1 length as well as skull length ( Meloro et al . 2007 ) . Lynx spelaeus ( Boule , 1906 ) The cave lynx is an enigmatic form that in the Italian middle Pleistocene is represented by theremains of Valdemino ( Sala et al . 1992 ) . What is not clear , is the evolution of such form and itsrelationship with the actual lynxes . Ficcarelli and Torre ( 1977 ) proposed that L . issiodorensis isancestral to the modern Spanish lynx ( Lynx pardinus ) and suggested for Pleistocene specimens a sub - specific rank ( L . pardina spelea ) for which an Asian immigrations seems likely . Recently , Testu ( 2006 ) described specimens of L . spelaeus from French localities ( Caune de L ’ Arago ; L ’ Obervatoire ) . These forms differ slightly in morphology from L . pardinus as well as L . lynx : theupper carnassial is larger while P 3 is more elongated than larger ; in the lower dentition the m 1 morphology is quite distinctive with the relative presence of metaconid - talonid complex in L . spelaeus while this feature lack in L . pardinus . Long bones morphology is generally robust and less elongated than in modern lynx . The dentition traits of L . spelaeus sometimes exhibit overlap with L . issiodorensis or L . lynx and this fact suggests that a gradual evolution occurred in Eurasiatic lynx starting from the Villafranchian Etouaires lynx , to an intermediate cave forms which splits inEuropean lynx ( L . lynx ) and endemic Spanish form ( L . pardinus ) ( cf . Testu 2006 ) . The Europeanlynx will be common in the Late Pleistocene all over Italian fossil record ( Rustioni et al . 1995 ) . The palaeoecology of cave lynx is unknown but its morphological features suggest a similarbehaviour to modern lynx . Body mass reconstruction based on m 1 length from Valdemino gives avalue of 23 kilograms only slightly large than Etouaires lynx ( Meloro et al . 2007 ) . Lynx issiodorensis Etouaires 스라소니 라 불리 는 이 녀석 은 체중 이 약 22 킬로 정도 로 추산 된다 하 며 . . 잘 발달 된 목부 위 와 그리고 아래 턱부위 를 보 면 강한 치 악력 을 지녔 다는 것 을 알 수 있 다 합니다 그리고 Lynx spelaeus , 동굴 스라소니 는 체중 이 23 킬로 로 추산 된다 합니다 . Panthera gombaszoegensis ( Kretzoi , 1938 ) This large cat is the oldest pantherine of Europe . The Italian fossil material is one the richest ( among the other Villafranchian European localities ) and it has been described by Del Campana ( 1916 ) and later by Ficcarelli and Torre ( 1979 b ) . Compared with the saber toothed cats , P . gombaszoegensis is characterized by less developed upper canine that are larger ; the dentition is generally not extremely reduced . Skull exhibits a frontal bone tiger like while the sagittal crest is strong as in the jaguar ( Panthera onca ) . Nasal process extends to the extreme of nasal bone ( Fig . 1 . 14 ) like in leopard ( P . pardus ) and jaguar . In the upper dentition the P 3 ( even if with littlediagnostic value ) shows anterior and posterior accessory cusps , P 4 has a distinctive cingulum , the canine is robust and incisor battery are arranged in a straight line with the third being larger ( O ’ Regan and Turner 2004 ) . In the mandible the lower canine is large and conical , the p 4 has alarge protocone together with anterior and posterior cusps , the lower carnassial is much distinctive with protoconid longer than paraconid and with rudimentary metaconid present . Long bones anatomy resembles some traits typical of the jaguar : the tibia is robust and the digital cavity of femur is similar to P . onca even if the neck of the femur is much more developed than in P . pardus and jaguar . In the fore limb the humerus shows several traits common to the lion P . leo ( e . g condyle ) while the trochlea is similar in size to the jaguar . Instead , the radius is much more similar to the tiger ( P . tigris ) . The oleocran region of the ulna is more developed than in the leopard . Metapodials are generally robust and jaguar like ( Del Campana 1916 ) . Panthera gombaszoegensis , 유럽 재규어 라 불리 는 이 녀석 은 90 ~ 210 킬로 에 달한다 말 해 지 고 있 으며 이 자료 의 저자 는 이탈리아 의 표본 들 의 무게 로 추산 해 낸 무게 는 90 킬로 를 추산 했 군요 . Panthera leo ( Linnaeus , 1758 ) The lion is an extant species whose geographic range is today restricted to Africa and the Indian forest of Gir ( Barnett et al . 2006 ) . Its occurrence in European fossil record begins in Galerian localities ( Kurtén 1968 ) . The oldest record is for Isernia La Pineta ( Sala 1990 ) that recently has been re - dated at 600 - 500 ka ( Coltorti et al . 2005 ) . in PCOM Galerian 3 , the lion is recorded at Fontana Ranuccio and in the Aurelian this species still to be rare at Torre in Pietra and Castel di Guido ( Capasso Barbato and Minieri 1987 ; Palombo et al . 2003 ; Raia et al . 2006 b ) . The Pleistocene lion is generally subdivided in two sub species : the ancient form P . leo fossilis of Middle Pleistocene and the Upper Pleistocene P . leo spelaea that is less massive and achieve similar upper size limits of extant lions ( Kurtén 1968 ) . Recently , Sotnikova and Nikolskiy ( 2006 ) proposed a specific rank for the Upper Pleistocene form ( P . spelaea ) on the basis of several skull and upper carnassial traits . Although this hypothesis cannot be excluded , I prefer to use aconservative approach by excluding sub - specific or new specificic denomination for Pleistocene lion . In keeping with Testu ( 2006 ) it is possible to consider the form fossilis and spelaea as chrono morphotypes whose morphological variability is the reflection of different climate . In such perspective the Italian specimens of middle Pleistocene lion are representative of the P . leo fossilis morphotype . This form is characterised by being larger and more robust than Upper Pleistocene and extant lion . Teeth are massive and less compressed . A shorter metacone can be discriminated in the upper cheek tooth . In m 1 protoconid is higher than paraconid , talonid is marked and a neat metaconid developed . The lower carnassial is generally more elongated than actual lion but less than in Upper Pleistocene lion . For middle Pleistocene there are non - complete skulls ( on the basis of Italian fossil record ) but Sotnikova and Nikolskiy ( 2006 ) describes features belonging to specimens from Azé ( French ) : the skull shares similar features with extant lion and Upper Pleistocene lion in being very large and with developed nuchal crest , the incisors are relatively smaller than in other lions , the orbits are small as well while the nuchal surface is broader . In the postcranial elements the humerus is massive with a medial epicondyle well developed on the supracondylar foramen ( present in felids and mustelids ) , metacarpal is massive with an elevated distal portion ( typical for felids ) . The femur is elongated and massive and the hindlimb is longer or similar in proportion to fore limb . The general morphological modification of Pleistocene lions appears to be a by product of their larger size hence a consequence of different environmental conditions . The palaeoecology of Pleistocene lion is generally based on several generalization that can be done by looking at the ecology of extant lion ( Schaller 1972 ) . The lion is a remarkable , highly dimorphic , opportunistic predator : its social behaviour is unique among felids ( usually lives in a pride familiar nucleus with multiple females and a dominant male ) and its prey preference is represented by medium - large ungulates ( in Serengeti the buffalo Syncerus caffer , the wildebeest , zebra ) . Such a preference causes overlap with several predators like spotted hyenas , African wild dogs but also cheetah . The latter two species evolved distinct hunting behaviour in order to avoid lion that is generally dominant by virtue of its larger size . Sometimes lion is capable to kill also very large ungulates like rhinoceros orjuvenile of elephants but they never constitute an important proportion in the diet ( Schaller 1972 ) . It has been suggested that the steppe bison was , probably , an important prey for the Pleistocene lion but such observation is valid on the basis of the bison “ Blue babe ” from a cave in Alaska ( Turner and Antón 1992 ) . For European lion there are no evidence of particular prey selection even if at Isernia La Pineta , for instance , the steppe bison ( Bison schoetensacki ) bones are really high in number but there are sign of human activity on them ( Anconetani and Peretto 1996 ) . It is not excluded that auroch ( Bos primigenius ) , wild boar ( Sus scropha ) and red deer ( Cervus elaphus ) could have been important for the middle Pleistocene European lion like gaur ( Bos gaurus ) , wild boar and Sambar deer ( Cervus unicolor ) are important for the tiger ( P . tigris ) ( Sunquist and Sunqist 1989 ; Kawanishi 2002 ) . It is noteworthy that some authors suggest several morphological similarities between Pleistocene lion and the tiger such that it can be considered more related to the latter . Recently , Yamaguchi et al . ( 2004 ) evidenced that the Pleistocene lion already lived in group but the typical mane ( now present in males as a secondary sexual trait ) was not present . For body mass I use an estimate of 183 kilograms as mean between males and females ( Meloro et al . 2007 ) . Panthera pardus ( Linnaeus , 1758 ) The leopard is a rare species in the Italian fossil record . Kotsakis and Palombo ( 1979 ) reviewed its distribution in the Italian fossil record . For the middle Pleistocene the earliest record is Valdemino ( Galerian 2 ) , than in the Aurelian the leopard is recorded at Capri and Prati Fiscali ( Palombo et al . 2003 ; Raia et al . 2006 b ) . Its distribution will be broader in the Upper Pleistocene , and in somelocalities leopard remains were high in number ( 100 individuals in Caverna degli Equi DelCampana 1947 ) . The skull of this predator exhibits a short muzzle with a more developedpostorbital region than in the jaguar ( Fig . 1 . 15 ) ; there are two typical inflations ( frontal and parietal region ) and one is below the alveolus of the upper canine ; the orbits are circular and the bullae are elongated . Upper premolars are variable in number and the P 1 can be supported by supranumerary aleveolus ; upper canines are flat in lingual side and the cheek tooth P 4 is compressed . In the mandible , the profile is generally not curved with a long diastema between canine and p 3 and also a short diastema can be present between p 3 and p 4 . The massetteric fossa is well developed and can achieve in length the posterior extremity of m 1 suggesting a strong masticatory power . The premolars do not exhibit significant morphologies ( in p 3 the anterior cusp can disappear while p 4 show tri - cusps ) , the m 1 is typical in its hypercarnivore morphology without metaconid and the protoconid is generally higher than paraconid ( cf . Testu 2006 ) . On the basis of Upper Pleistocene fossils , Del Campana ( 1947 ) demonstrated that no significantdifferences can be found in the morphology of the post - cranial anatomy of Pleistocene leopard and the extant one even if the Equi specimens exhibited a high proportion of arthritis in humerus . The hind limb is proportionally longer than the forelimb ( Lewis 1997 ) and this confers a highadaptability in tree climbing . The leopard is considered one of the most peculiar biotic agents in the taphonomy of several caves especially in Africa because of its capacity to drag carcasses on trees or near caves : among the other prey also hominid were founds in Swartkans ( Turner and Antón 1997 ) . The extant leopard is sexually dimorphic and can achieve 70 kilograms ( males ) : for the Pleisotcene form I use an estimate of 60 kilograms ( Meloro et al . 2007 ) . Panthera pardus , 유럽 표범 이 라 불리 는 이 녀석 은 수컷 이 70 킬로 에 달할 수 있 다 하 며 저자 는 빙하기 의 유럽 표범 들 은 60 킬로 정도 라 추산 했 군요 . 마지막 으로 Panthera leo ( spelaea ) 즉 동굴 사자 는 갈기 가 별로 없 었 다 하 며 체중 은 암수 평균 을 183 킬로 로 추산 한다 합니다 . 그렇 다면 수컷 의 체중 은 , ,? 한 210 ~ 220 킬로 정도 라 말 하 겠 군요 . 현생 사자 보다 20 % 가량 큰 수치 정도 . . 입니다 . 예전 에 는 동굴사자 가 아메리카사자 보다 더 크 다는 말 이 있 었 으나 요새 는 말 이 바뀌 어서 아메리카 사자 가 가장 큰 사자 이 며 동굴사자 는 생각 만큼 그렇게 크 진 않 았 다고 하 는 듯 합니다 . 위 에서부터 순서 대로 아메리카사자 - 동굴사자 - 아프 라 키 사자 의 두개골 입니다 . 동굴사자 와 아프리카 사자 의 차이 는 그렇게 크 진 않 지만 아메리카사자 와 의 차이 는 상당 하 군요 . 이게 지금 저 셋 의 크기 를 보여 주 는 비교 도 라고 생각 이 드 는군요 .