Megantereon Whitei Research Articles

On the ecological scenario of the first hominin dispersal out of Africa

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 100kg 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.

Zoogeographic significance of Dmanisi large mammal assemblage

We undertake a comparative mammalian zoogeographic analysis with the aim of revealing the extent to which the Dmanisi Early Pleistocene large mammal assemblage resembles, at the genus level, African, Arabian, and Eurasian localities of similar age. The inclusion of Old World Pliocene and Pleistocene mammalian faunas provides us with insights into the provincial origins of specific mammalian taxa and permits us to assess the relative affiliation of the Dmanisi mammalian faunas to other faunas in the Old World. Our analysis also allows us to consider hypotheses about the timing and direction of zoogeographic connections between western Eurasia and Africa during the Early Pleistocene. We utilize multiple zoogeographic analytical tools as a cross-comparison of Dmanisi with 42 other Eurasian and African mammalian-bearing localities between 2.7 and 0.7 Ma. Overall, we find that Dmanisi compares most closely with a subgroup of Greek, Italian, and Spanish localities that are slightly younger than Dmanisi itself. This could suggest a progressive dispersal from East to West of the large mammal communities during the late Early Pleistocene and the first occurrence at Dmanisi, and then later in Western Europe, of some taxa such as Stephanorhinus ex gr. etruscus-hundsheimensis, Equus altidens, Bison georgicus, Soergelia minor, Megantereon whitei, Canis borjgali, Canis (Xenocyon) lycaonoides. Dmanisi's habitats included drier areas, probably of open wooded savannah and grassland and by mountainous to semiarid rocky terrain. There is evidence that Dmanisi records short intervals of increased aridity in the middle part of the succession contemporaneous with the occurrence of Homo.

A comparative study of the Early Pleistocene carnivore guild from Dmanisi (Georgia)

The carnivore guild of the Early Pleistocene site of Dmanisi is among the most diverse of the Early Pleistocene of the entire Old World. It includes 14 carnivoran taxa: Homotherium latidens, Megantereon whitei, Panthera onca georgica, Acinonyx pardinensis, Lynx issiodorensis; Pachycrocuta brevirostris; Canis (Xenocyon) lycaonoides, Canis borjgali, Vulpes alopecoides; Ursus etruscus; Lutra sp., Martes sp., Meles sp., and Pannonictis sp. The analysis of this rich carnivore guild was carried out under different methodological approaches to compare the assemblage with other chronological coeval European, Asian, and African sites from a paleobiological perspective. To achieve the goal, we used a permutational hierarchical method called boostrapping cluster analysis based on taxonomic absence/presence matrices (at both generic and specific level) and on ecological matrices (considering dietary preferences/hunting strategies of each carnivoran) and carried out Mantels tests assessing magnitude of time, space, ecology, and taxonomy as source of difference between guilds. Our results suggest a close similarity among the Dmanisi carnivore assemblage and other guilds recorded from European late Villafranchian sites such as Pirro Nord, Venta Micena, and Apollonia 1 and, in a lesser extent, to European Epivillafranchian sites as Vallonnet, Untermassfeld, or the Vallparadís Section. Early to Middle Pleistocene Asian carnivore assemblages display several similarities with the Dmanisi guild mainly in the record and diversity of felid and the canid ecomorphotypes. Eastern African sites such as Olduvai and Omo, as well as South African sites, display a lower similarity with the studied sample, basically for the most diverse hyenid taphocoenoses. To sum up, the present study suggests a close similarity between the Dmanisi carnivore guild and other European Late Early Pleistocene assemblages without close parallels with African or Asian assemblages.

Megantereon (Carnivora, Felidae) in the late Early Pleistocene in China and its implications for paleobiogeography

Specimens of the genus Megantereon from younger strata (late Early Pleistocene to Middle Pleistocene) in China are reviewed in the presented work. The specimens found in Linyi, Shanxi Province, which were previously identified as Megantereon nihowanensis, have been reexamined and their identity has been revised to be Megantereon inexpectatus. With respect to the new specimens of M. inexpectatus found in the Jinyuan Cave, Dalian, Liaoning Province, the identification of the mandible specimen from the Gongwangling Homo site, which was named as a new species, Megantereon lantianensis, has been revised to M. inexpectatus. Known records of M. inexpectatus have been reported in Linyi, Shanxi, and Lantian, Shaanxi Province, in older deposits from the late Early Pleistocene, and younger deposits from the Middle Pleistocene in Zhoukoudian, Beijing. In morphological comparisons and phylogenetic discussions, the best candidate origin of M. inexpectatus is Africa, where a relatively advanced lineage gave rise to M. inexpectatus and its relative Megantereon whitei. A significant dispersal event of M. inexpectatus occurred during the late Early Pleistocene, represented by the immigration of M. whitei from Africa to Europe and the occurrence of M. inexpectatus in Asia. There is no evidence indicating that this biotic event was initiated by environmental change, with survival pressure from hominins likely to have been the main factor that forced Megantereon to migrate into Eurasia.

On a New Find of Megantereon (Carnivora, Felidae, Machairodontinae) from the Early Pleistocene of Trlica (Montenegro, the Central Balkans)

A canine of Megantereon whitei (Broom, 1937), a large representative of the genus Megantereon from Trlica Cave in Montenegro, a multilayered mammal locality with traces of hominine activity, is described. The specimen was found in Bed 9 of the section during excavations in 2010–2014. This record reflects the first penetration of the African species M. whitei into the Balkans. The mammal fauna from Trlica (TRL11–TRL10, 1.8–1.5 Ma) includes the Eurasian species M. cultridens, represented by a canine found in Bed 10.

The new Carnivore remains from the Early Pleistocene Yanliang Gigantopithecus fauna, Guangxi, South China

Abstract New Carnivore remains associated with Gigantopithecus blacki are described from Yanliang Cave, Chongzuo, Guangxi Zhuang Autonomous Region, South China. New specimens of Megantereon have been identified as Megantereon microta sp. nov. based on the previously unknown characteristics, such as small overall size, p4 and m1 dental proportions, p3 with two separated roots and large P4 protocone. M. microta sp. nov. differs from Megantereon whitei from Africa mainly by the characters of p3 and lacking a p3–p4 diastema. Compared with other Megantereon species from China, M. microta sp. nov. is considered to be more primitive in dental characters, such as large P4 protocone, p3 with two separated roots, and short canine-p3 diastema. The Pachycrocuta licenti and Ursus thibetanus from Yanliang Cave are consistent with their appearance during the early Early Pleistocene in southern China. Combining the biostratigraphical and biochronological evidence, the Yanliang fauna can be best correlated with Baikong fauna in the same area, which is the early Early Pleistocene (∼2.0 Ma).

CT scanning analysis of Megantereon whitei (Carnivora, Machairodontinae) from Monte Argentario (Early Pleistocene, central Italy): evidence of atavistic teeth.

CT scanning analysis applied to vertebrate palaeontology is providing an increasing number of data of great interest. This method can be used in many branches of palaeontology such as the investigation of all the fossilized elements in a hard matrix and the hidden structures in the bones. A large number of pathologies are "hidden", completely or partially invisible on the external surface of the bones because their development took place within the bones. However, the study of these diseases and abnormalities plays a crucial role in our understanding of evolutionary and adaptive processes of extinct taxa. The analysis of a partial skeleton of the sabre-toothed felid Megantereon whitei from the Early Pleistocene karst filling deposits of Monte Argentario (Tuscany, Italy) has been carried out. The CT scanning analysis put in evidence the presence of supernumerary teeth (P(2)) and the absence of P3 in the mandible. The occurrence of P(2) can be considered as an evidence of atavism. Such an archaic feature is recorded for the first time in Megantereon.

A new record of the saber-toothed cat Megantereon (Felidae, Machairodontinae) from an Early Pleistocene Gigantopithecus fauna, Yanliang Cave, Fusui, Guangxi, South China

Here, we describe saber-toothed cat remains from the early Early Pleistocene (∼2.0 Ma) of Yanliang Cave, Fusui County, Guangxi Province, South China. Specimens, including a complete left dentary and right m1, are identified as Megantereon based on mandibular and dental characters. The Yanliang Megantereon is the smallest representative of the genus from Asia. Overall size, p4 and m1 dental proportions, the disposition of the p3, and dentary measurements align it with characters attributed to Megantereon whitei, a taxon not recorded from eastern Asia or China. Megantereon whitei is thought to be the most derived member of the genus. If the remains are M. whitei, or a closely related form, this represents an early occurrence of the group outside Africa. Other Megantereon specimens from China, that are roughly the same geologic age, or even younger, are considered to have a more primitive dentition than M. whitei. The Yanliang Megantereon, and our review, further demonstrates uncertainty in the evolutionary history of the group, the overall need for tighter chronologic control, and a systematic revision of the genus that includes discrete morphological characters.

First occurrence of Soergelia (Ovibovini, Bovidae, Mammalia) in the Early Pleistocene of Italy

The revision of the Early Pleistocene collection from the site of Monte Argentario (southern Tuscany, Italy), housed at the Istituto Italiano di Paleontologia Umana (Roma), revealed the occurrence of a mostly complete left horn core, attributable to a small-middle size bovid species corresponding to the genus of Asian origin Soergelia (Soergelia sp. cf. Soergelia minor). This is the first finding of the genus in the Italian Peninsula. The Monte Argentario fauna associated with Soergelia includes the African origin saber-toothed tiger Megantereon whitei and other Late Villafranchian species.

Early human dispersals into the Iberian Peninsula: A comment on and

The carnivore guild of the Early Pleistocene site of Dmanisi is among the most diverse of the Early Pleistocene of the entire Old World. It includes 14 carnivoran taxa: Homotherium latidens, Megantereon whitei, Panthera onca georgica, Acinonyx pardinensis, Lynx issiodorensis; Pachycrocuta brevirostris; Canis (Xenocyon) lycaonoides, Canis borjgali, Vulpes alopecoides; Ursus etruscus; Lutra sp., Martes sp., Meles sp., and Pannonictis sp. The analysis of this rich carnivore guild was carried out under different methodological approaches to compare the assemblage with other chronological coeval European, Asian, and African sites from a paleobiological perspective. To achieve the goal, we used a permutational hierarchical method called boostrapping cluster analysis based on taxonomic absence/presence matrices (at both generic and specific level) and on ecological matrices (considering dietary preferences/hunting strategies of each carnivoran) and carried out Mantels tests assessing magnitude of time, space, ecology, and taxonomy as source of difference between guilds. Our results suggest a close similarity among the Dmanisi carnivore assemblage and other guilds recorded from European late Villafranchian sites such as Pirro Nord, Venta Micena, and Apollonia 1 and, in a lesser extent, to European Epivillafranchian sites as Vallonnet, Untermassfeld, or the Vallparadís Section. Early to Middle Pleistocene Asian carnivore assemblages display several similarities with the Dmanisi guild mainly in the record and diversity of felid and the canid ecomorphotypes. Eastern African sites such as Olduvai and Omo, as well as South African sites, display a lower similarity with the studied sample, basically for the most diverse hyenid taphocoenoses. To sum up, the present study suggests a close similarity between the Dmanisi carnivore guild and other European Late Early Pleistocene assemblages without close parallels with African or Asian assemblages.

The African species Megantereon whitei from the Early Pleistocene of Monte Argentario (South Tuscany, Central Italy)

A partial skull and articulated postcranial elements of Megantereon whitei have been recorded during the 1950s from a karst deposit in the Monte Argentario area (Grosseto). These fossils recently became available for study. The bones are quite well preserved, included in a hard reddish matrix with calcareous clasts. The fossil is part of a faunal assemblage referred to the Late Villafranchian (Early Pleistocene). M. whitei from Monte Argentario is characterised by elongated upper canines, stronger than those of the Upper Valdarno (Tuscany) specimens and similar to the Pirro Nord (Apulia) fossil. The manus is robust, the first phalanges are quite long in comparison to the metacarpals. The Early Pleistocene European M. whitei represents an African element that took part in the faunal dispersal from Africa to Europe that occurred at the Plio-Pleistocene transition.

Tracing the ecophysiology of ungulates and predator–prey relationships in an early Pleistocene large mammal community

Research into the reconstruction of ancient communities in terms of dietary regimes, habitat preferences and ecological interactions of species has focused predominantly on biogeochemistry or ecomorphology alone and not in combination. The Venta Micena site (Orce, Guadix–Baza basin, SE Spain) has an early Pleistocene vertebrate assemblage with exceptional biomolecular preservation. Collagen was successfully extracted from 77 bone samples of 18 species of large mammals, which allowed analyses of carbon- and nitrogen-isotopes. δ 13C, δ 15N and δ 18O ratios combined with ecomorphological indexes provide interesting clues on the autecology and palaeophysiology of extinct species, which help in deciphering aspects of community trophic structure and predator–prey interactions. Specifically, morphometric ratios (e.g., hypsodonty index and relative length of the lower premolar tooth row; [Palmqvist, P., Gröcke, D.R., Arribas, A., Fariña, R. 2003. Paleoecological reconstruction of a lower Pleistocene large mammals community using biogeochemical ( δ 13C, δ 15N, δ 18O, Sr:Zn) and ecomorphological approaches. Paleobiology 29, 204–228.]) allow classifying the ungulates among grazers ( Equus altidens, Bison sp., Praeovibos sp., Hemitragus albus, Hippopotamus antiquus, and Mammuthus meridionalis), mized-feeders ( Soergelia minor and Pseudodama sp.) and browsers ( Stephanorhinus sp. and Praemegaceros cf. verticornis). However, δ 13C values reveal that these ungulates consumed exclusively C 3 plants and significant differences in isotopic values between perissodactyls (monogastric, hindgut fermenters) and ruminants (foregut fermenters) must reflect physiological differences related to their rates of methane production and digestive efficiency. δ 18O ratios allow the interpretation of the dietary water source of these species, suggesting that fallow deer Pseudodama sp., goat H. albus and ovibovine S. minor obtained a significant fraction of their metabolic water from the vegetation consumed. Carnivore species have higher δ 15N values than herbivores, which records the isotopic enrichment expected with an increase in trophic level. However, the unexpectedly high δ 15N values of hippo H. antiquus and muskoxen Praeovibos sp. suggest that these ungulates predominantly consumed aquatic plants and lichens, respectively. Inferences on predator–prey relationships within this ancient community, derived from the dual linear mixing model, indicate resource partitioning among sympatric predators, suggesting that sabre-tooth Megantereon whitei and jaguar Panthera cf. gombaszoegensis were ambushers of forest environments while sabre-tooth Homotherium latidens and wild dog Lycaon lycaonoides were coursing predators in open habitat . The giant, short-faced hyena Pachycrocuta brevirostris scavenged the prey of these hypercarnivores.

Evidence of earliest human occurrence in Europe: the site of Pirro Nord (Southern Italy)

Some flint lithic artifacts were discovered in the fissure fillings of the well-known Pirro Nord site (Apulia, Southern Italy). The lithic industry, composed by three cores and some flakes, has been found to be associated to an Early Pleistocene vertebrate fossil assemblage. The fossil association contains a wide range of micromammals, including Allophaiomys ruffoi and Episoriculus gibberodon and large mammals including Bison degiulii and Equus altidens together with African elements as the gelada baboon Theropithecus and the saber-toothed cat Megantereon whitei. It defines the latest Villafranchian chronological unit (Pirro Nord Faunal Unit) in the Western European mammal biochronology. The lithic industry of Pirro Nord represents the oldest occurrence of the genus Homo in Europe as it is attributable to a chronological interval between 1.3 and 1.7 Ma. This supports the hypothesis that the genus Homo, with Oldowaian technology, extended its range in Europe, probably from western Asia, during the first half of the Early Pleistocene. The new discovery from Pirro Nord changes the chronology of the first arrival of hominids in Europe and offers new perspectives in the debate about the human dispersal in the Early Pleistocene.

Occurrence of Theropithecus sp. in the Late Villafranchian of Southern Italy and implication for Early Pleistocene “out of Africa” dispersals

Three cervical vertebrae of a large primate have been identified in the revision of the faunal assemblage collected during the late '70s from fissure fillings in the area known as Pirro Nord in southern Italy. By size the three vertebrae are comparable to the large fossil species Theropithecus oswaldi, however we prefer here to attribute the remains to Theropithecus sp. The Pirro Nord Theropithecus represents the second finding outside of Africa of this genus in association with Megantereon whitei, after the site of ‘Ubeidiya. The biochronology of Pirro Nord faunal assemblage demonstrates that the arrival into Europe of Theropithecus occurred earlier than thought: in the range of approx 1.6–1.3 Ma. The identification of an European “African assemblage” ( Theropithecus associated with Megantereon whitei) has consequences for interpretation of an “out of Africa” (genus Homo) event around the Plio-Pleistocene transition.

Paleoecological reconstruction of a lower Pleistocene large mammal community using biogeochemical (δ13C, δ15N, δ18O, Sr:Zn) and ecomorphological approaches

Ecomorphological and biogeochemical (trace element, and carbon, nitrogen, and oxygen isotope ratios) analyses have been used for determining the dietary niches and habitat preferences of large mammals from lower Pleistocene deposits at Venta Micena (Guadix-Baza Basin, Spain). The combination of these two approaches takes advantage of the strengths and overcome the weakness of both approaches. The range of δ13Ccollagen values for ungulate species indicates that C3 plants were dominant in the diet of these mammals. δ13Ccollagen values vary among ungulates: perissodactyls have the lowest values and bovids the highest ones, with cervids showing intermediate values. The hypsodonty index measured in lower molar teeth and the relative length of the lower premolar tooth row indicate that the horse, Equus altidens, was a grazing species, whereas the rhino, Stephanorhinus etruscus, was a mixed feeder in open habitats. The similar δ13Ccollagen values shown in both perissodactyls does not reflect differences in feeding behavior with other ungulates, but rather a lower isotope enrichment factor in these monogastric herbivores than in ruminants, owing to their lower metabolic efficiency. The cervids Eucladoceros giulii and Dama sp. show low hypsodonty values, indicating that they were mixed feeders or browsers from forested habitats, an ecomorphologically based conclusion corroborated in the former by its low δ15Ncollagen content (canopy effect). Bovid species (Bovini aff. Leptobos, Soergelia minor, and Hemitragus albus) presumably inhabited open environments, according to their comparatively high hypsodonty and δ15Ncollagen values. Carnivore species (Homotherium latidens, Megantereon whitei, Pachycrocuta brevirostris, Canis falconeri, and Canis etruscus) exhibit higher δ15Ncollagen values than ungulates. These results record the isotopic enrichment expected with an increase in trophic level and are also supported by low bone Sr.Zn ratios. The elevated δ15Ncollagen value for a sample of Mammuthus meridionalis, which came from an individual with unfused epiphyses, confirms that it was a suckling animal. The δ15Ncollagen value of the scimitar-cat H. latidens is well above that obtained for the young individual of Mammuthus, which indicates that juvenile elephants were an important part of its diet. The hippo, Hippopotamus antiquus, yielded unexpectedly high δ15Ncollagen values, which suggest feeding on aquatic, non-N2-fixing plants. The high δ18Ohydroxyl values of bovids Hemitragus and Soergelia and of cervid Dama indicate that these ungulates obtained most of their water requirements from the vegetation. The megaherbivores and Eucladoceros exhibit the lowest δ18Ohydroxyl values, which suggest increased water dependence for them. Paleosynecological analysis was based on the relative abundance of species of large mammals from different ecological categories, determined by feeding behavior and locomotion types. The comparison of the frequencies of such categories in Venta Micena with those found in modern African communities indicates that the composition of the paleocommunity closely resembles those of savannas with tall grass and shrubs. The net above-ground primary productivity estimated for the on-crop biomass of the mammalian species preserved in the fossil assemblage also yields a figure congruent with that expected for an open environment.

On the presence of Megantereon whitei at the South Turkwel Hominid Site, northern Kenya

Werdelin and Lewis (2000) recently described a new species of the saber-tooth genus Megantereon (Mammalia, Carnivora, Felidae) from Late Pliocene (∼3.5 Ma) deposits of the Nachukui Formation south of the Turkwel River in Turkana, northern Kenya (Ward et al., 1999). The new species, Megantereon ekidoit, is said to represent the earliest occurrence of this genus in Africa. It was described from a right mandibular ramus (KNM-St 23812) with i3–ml. As discussed below, the post-canine diastema is longer than the cheek-tooth row, since there does not appear to have been a third premolar (p3), and the coronoid process is relatively well developed, showing a hook-shaped morphology (Werdelin and Lewis, 2000: Fig. 1). The fourth premolar (p4) and the lower carnassial tooth (ml) show considerable wear.

Prey selection by terrestrial carnivores in a lower Pleistocene paleocommunity

We report quantitative paleoecologic data on the large mammal assemblage preserved in lower Pleistocene deposits at Venta Micena (Orce, Granada, southeastern Spain). Taphonomic studies show that bones were collected mainly by hyaenids, which transported and deposited them near shallow dens. Differential fragmentation of major long bones was produced by hyaenas as a function of their density and marrow content. Strong selection of prey by carnivores—which preferentially killed juveniles, females, and individuals with diminished locomotor capabilities among ungulate prey species of larger body size—is indicated by (1) the abundance of remains of juvenile ungulates in relation to the average weight of adult individuals in each species, (2) attritional mortality profiles for ungulate species deduced from crown height measurements, (3) the presence of many metapodials with different osteopathologies in their epiphyses, such as arthrosis, and (4) a biased intersexual ratio of large bovids. Comparison of the frequencies with which modern African carnivores kill and scavenge ungulates from various size classes with the abundance of these size categories in the assemblage suggests that the Venta Micena hyaena (Pachycrocuta brevirostris) was a bone-cracking scavenger that fed largely on carcasses of ungulates preyed upon and partially consumed by fresh meat-eating carnivores such us saber-toothed felids (Homotherium latidens and Megantereon whitei) and wild dogs (Canis falconeri).

Presence of the African Machairodont Megantereon whitei(Broom, 1937) (Felidae, Carnivora, Mammalia) in the Lower Pleistocene Site of Venta Micena (Orce, Granada, Spain), with some Considerations on the Origin, Evolution and Dispersal of the Genus

We have made a comparative analysis of dental morphometries of the Megantereonspecimens found in the Lower Pleistocene site at Venta Micena (Orce, Granada, Southeastern Spain) with representatives of the genus throughout the Old and New World. The methodologies used in the study include discriminant functions and principal component analysis. The results obtained point to the presence of three distinct species of this genus: M. cultridens(Cuvier), indigenous to North America from the Lower Pliocene onwards, which in the Indian subcontinent gave rise to the larger sized species M. falconeriPomel during the Upper Pliocene. M. cultridenssurvived until the Middle Pleistocene in China, and its African descendant M. whiteiBroom colonized Europe at the time of the Plio-Pleistocene boundary, reaching Dmanisi via the Near East and southern part of the Iberian Peninsula at Venta Micena via the Strait of Gibraltar.