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Phylogeny of the giant deer with palmate brow tines Megaloceros from west and Sinomegaceros from east Eurasia
Abstract
Giant deer with palmate brow tines were placed either in a single genus, or those from west Eurasia were placed in Megaloceros and those from the east in Sinomegaceros, which implies local evolution. In order to see whether dispersals between the two areas may have occurred, we studied the morphology and interpreted the phylogeny of these deer.The phylogenetic model proposed does not support any dispersals between east and west after the appearance of the first species of each genus, but it does support the recognition of two separate genera.The ecological significance of some of the morphological characters is interpreted. Thicker molar enamel, reduced posterior molars, larger premolars and a P4 with a lesser degree of “molarisation” suggests that Megaloceros savini adapted to a diet that includes coarser and harder food. Elevated mandibular condyles, thin molar enamel, and P4 that are predominantly “molarized” suggest that Megaloceros giganteus and Sinomegaceros yabei shifted towards grazing larger quantities of not very hard food. Very robust metapodials evolved several times in western Eurasian giant deer: during the Early Pleistocene in the Megaloceros solilhacus group, not later than the late Middle Pleistocene in M. giganteus and it increased in the Late Pleistocene M. giganteus. Since the character did not evolve in the east Eurasian giant deer, it might be related to some aspect of the west Eurasian environment, that was not present in east Eurasia.
... 3 In contrast to the controversial status of giant deer taxonomy in pre-Pleistocene, and even into the Early and Middle Pleistocene in Europe, researchers agree on a morphology-based division between the Late Pleistocene Megaloceros giganteus of western Eurasia and an array of Pleistocene Sinomegaceros species from eastern Eurasia (Figure 1). 3,6 This is based, among other features, on the extraordinarily broad and flattened brow (basal) tine, expanded in most Sinomegaceros species beyond the more modest flattening seen in Megaloceros and unique among living and fossil deer ( Figure 2). Up to nine species of Sinomegaceros have been proposed, from Central Asia, China, and Japan 6 (five were recognized by Vislobokova 3 ). ...
... Analysis from morphological characters suggested that the eastern Eurasia cluster, Sinomegaceros, diverged from Megaloceros more than 1 Ma. 3,6 Fossil records revealed that the range of Sinomegaceros covered continental areas from approximately 50 to 25 N, mainly in present-day eastern and northeastern China and the Japanese islands. 6,12 In China, there are several well-known fossil species of Sinomegaceros from different stages of the Pleistocene, with Sinomegaceros konwanlinensis, S. pachyosteus, and S. ordosianus representing the Early, Middle, and Late Pleistocene, respectively. ...
... 3,6 Fossil records revealed that the range of Sinomegaceros covered continental areas from approximately 50 to 25 N, mainly in present-day eastern and northeastern China and the Japanese islands. 6,12 In China, there are several well-known fossil species of Sinomegaceros from different stages of the Pleistocene, with Sinomegaceros konwanlinensis, S. pachyosteus, and S. ordosianus representing the Early, Middle, and Late Pleistocene, respectively. 13 In Japan, Sinomegaceros yabei appeared from the second part of the Middle Pleistocene to the Pleistocene/Holocene boundary. ...
The giant deer, widespread in northern Eurasia during the Late Pleistocene, have been classified as western Megaloceros and eastern Sinomegaceros through morphological studies. While Megaloceros's evolutionary history has been unveiled through mitogenomes, Sinomegaceros remains molecularly unexplored. Herein, we generated mitogenomes of giant deer from East Asia. We find that, in contrast to the morphological differences between Megaloceros and Sinomegaceros, they are mixed in the mitochondrial phylogeny, and Siberian specimens suggest a range contact or overlap between these two groups. Meanwhile, one deep divergent clade and another surviving until 20.1 thousand years ago (Ka) were detected in northeastern China, the latter implying this area as a potential refugium during the Last Glacial Maximum (LGM). Moreover, stable isotope analyses indicate correlations between climate-introduced vegetation changes and giant deer extinction. Our study demonstrates the genetic relationship between eastern and western giant deer and explores the promoters of their extirpation in northern East Asia.
... 周家油坊 Zhoujiayoufang [36] 闫家岗 Yanjiagang [33] 灵井 Lingjing [37] 周口店 ZKD 1 [11] 周口店 ZKD 13 [5] 陕西洛川 [26] Luochuan [14] 和肿骨鹿 (S. pachyosteus) [ [5,21,22] 和眉枝形态 [3,4] 经常被作为大角鹿系统分类的重要特征，不过下颌骨是否可以作为分类的重要依据尚存争 议，此外关于大角鹿下颌骨肿厚指数的计算，存在若干不同方法 [23] 。Azzaroli 根据眉枝形 态将大角鹿类分成两组：一组以爱尔兰大角鹿为代表，眉枝呈掌状；另一组以直角大角鹿 (Megaloceros verticornis) 为代表，眉枝横截面为圆形 [3] 。从掌状眉枝来看，其所在平面与 头骨矢状面之间的关系是区分不同属种的一个重要特征，大致可以分为三种类型：垂直于 头骨矢状面、与头骨矢状面斜交、平行于头骨矢状面 [4] 。此外，van der Made 和 Tong [4] 将 ...
... 周家油坊 Zhoujiayoufang [36] 闫家岗 Yanjiagang [33] 灵井 Lingjing [37] 周口店 ZKD 1 [11] 周口店 ZKD 13 [5] 陕西洛川 [26] Luochuan [14] 和肿骨鹿 (S. pachyosteus) [ [5,21,22] 和眉枝形态 [3,4] 经常被作为大角鹿系统分类的重要特征，不过下颌骨是否可以作为分类的重要依据尚存争 议，此外关于大角鹿下颌骨肿厚指数的计算，存在若干不同方法 [23] 。Azzaroli 根据眉枝形 态将大角鹿类分成两组：一组以爱尔兰大角鹿为代表，眉枝呈掌状；另一组以直角大角鹿 (Megaloceros verticornis) 为代表，眉枝横截面为圆形 [3] 。从掌状眉枝来看，其所在平面与 头骨矢状面之间的关系是区分不同属种的一个重要特征，大致可以分为三种类型：垂直于 头骨矢状面、与头骨矢状面斜交、平行于头骨矢状面 [4] 。此外，van der Made 和 Tong [4] 将 ...
... 周家油坊 Zhoujiayoufang [36] 闫家岗 Yanjiagang [33] 灵井 Lingjing [37] 周口店 ZKD 1 [11] 周口店 ZKD 13 [5] 陕西洛川 [26] Luochuan [14] 和肿骨鹿 (S. pachyosteus) [ [5,21,22] 和眉枝形态 [3,4] 经常被作为大角鹿系统分类的重要特征，不过下颌骨是否可以作为分类的重要依据尚存争 议，此外关于大角鹿下颌骨肿厚指数的计算，存在若干不同方法 [23] 。Azzaroli 根据眉枝形 态将大角鹿类分成两组：一组以爱尔兰大角鹿为代表，眉枝呈掌状；另一组以直角大角鹿 (Megaloceros verticornis) 为代表，眉枝横截面为圆形 [3] 。从掌状眉枝来看，其所在平面与 头骨矢状面之间的关系是区分不同属种的一个重要特征，大致可以分为三种类型：垂直于 头骨矢状面、与头骨矢状面斜交、平行于头骨矢状面 [4] 。此外，van der Made 和 Tong [4] 将 ...
Giant deer is among the most common animals of Mid-Late Pleistocene sites in northern China, and was one of the representative icons of the Pleistocene fauna in northern China, while the fossil materials are fairly poor and very few complete antlers were ever recovered, except those from Zhoukoudian site; on the other hand, the knowledge about the agerelated changes of the antlers is absolutely insufficient. Therefore, the previous taxonomic work based only on the antler features is open to questions.
In 2018, a quite complete antler of giant deer was recovered from the Nanbaishan site of Mesolithic age in Yüxian County, Hebei Province. The antler is quite big; brow tine and the palmation of beam are thin and fan-like, without palm tines; the brow tine and the palmation of beam run along two nearly parallel planes, but are not exactly parallel; the shaft of the beam bends at the basal part, but the sigmoid form is not prominent. In general morphology, the new antler is very close to that of Sinomegaceros ordosianus. The new specimen represents the most complete antler of S. ordosianus ever recovered. The dimensions (L×W) of the palmate is 670 × 526.8 mm; the dimensions (L×W) of the brow tine is 510 × 480 mm, the length and
circumference of the beam are 270 and 193 mm respectively, the circumference of the burr is 310 mm. The origin of the fossil-bearing strata is not clear yet, while its luminescence age is ca. 110 kyrs. The first phalange is robust, its greatest length is 77.7 mm; proximal width is 29.2 mm, transverse and antero-posterior diameters are 24.0 and 26.0 mm respectively.
In China, quite a number of megalocerine taxa had been named at the species and subspecies levels, namely Sinomegaceros pachyosteus, S. ordosianus, S. flabellatus, S. konwanlinensis, S. youngi, S. luochuanensis, S. sangganhoensis, S. o. mentougouensis and S. baotouensis, among which S. ordosianus is the most widely distributed species and has the richest fossil records. On the contrary, S. baotouensis is the least known species which only represented by one shed antler
and one metatarsal bone. With only a few exceptions, e.g. Tangshan near Nanjing, Zhoushan island in Zhejiang and Hualong Cave in Anhui, all the other megalocerine fossil sites in China occur north of the Yangtze River, and most of them are located in northern China. The fossils of S. ordosianus were frequently appeared in the prehistoric site, which indicates that the giant deer was very probably among the food sources of early humans.
... The acronyms of the collections where those specimens were studied are indicated in Table 5. The methods of study of the Cervidae and the way of measuring (Table 6) are similar to those used by Van der Made and Tong (2008). The measurements in the figures and in Tables 7 and 8 are in millimeters. ...
... We can observe that there is a tendency for the successive species to have the brow tine in progressively lower positions. This is a common evolutionary tendency in the Cervinae and has been documented for Dama (Van der Made 1999; Van der Made et al. 2016Made et al. , 2017, Megaloceros, and Sinomegaceros (Van der Made & Tong 2008;Van der Made 2015 and is also known from Megaceroides (Praemegaceros), while in Eucladoceros there is no clear tendency, but the different species differ in this feature (Van der Made & Dimitrijevic 2015). Though there are fewer data on the position of the middle tine (B) than on the brow tine (A), in successive samples or species, it also tends to originate in a progressively lower position. ...
... Though there are fewer data on the position of the middle tine (B) than on the brow tine (A), in successive samples or species, it also tends to originate in a progressively lower position. This is documented in Megaloceros (Van der Made & Tong 2008), but it occurs also in Dama. The positions in which the different tines originate from the main beam are not independent but are related: if the brow tine originates lower, the middle tine also originates lower (compare Figs. 8 and 10). ...
We describe fossils of a new species of fallow deer, Dama celiae. It is the end member of the lineage Dama farnetensis–D. vallonnetensis–D. roberti–D. celiae, which reduced the number of points of the antler from four to two, while the parallel lineage leading to the living fallow deer evolved more complex and palmate antlers. The fossils are from localities Pedro Jaro I and Orcasitas in the + 25–30-m terrace of the Manzanares river, which is correlated to MIS9 (337–300 ka) and which also yielded fossils of Megaloceros matritensis, a recently named species, end member of a lineage that survived longer than previously believed. A younger terrace of the Manzanares yielded remains of Haploidoceros, a rare deer known from two older localities in southern France and one younger locality in Spain. So many rare deer species in this valley indicates either endemism and a very special environment or that the record of fossil deer is much less known than generally assumed. Until recently, the European Middle Pleistocene record of deer had only one middle-sized species at a time. Now, it appears that there were up to three contemporaneous species of the size of a fallow deer. Acheulean lithic assemblages have been documented from the same sites as Dama celiae. This species was contemporaneous to Neanderthals with Acheulean culture. Cut marks suggest that it was consumed by them and probably was hunted
... The dental morphology is generally unspecialized, with primitive unmolarized P4 and occasional presence of small lingual cingulum in upper molars [89]. The phylogenetic relationship between Praedama and Megaloceros was accepted by van der Made and Tong [90], however, the cited authors doubted that P. savini could be a direct ancestral form of M. giganteus because of the divergence in mastication adaptations. Vislobokova [23,26,91] regards Praedama as a side phylogenetic branch of the Arvernoceros-Megaloceros lineage. ...
... The dental morphology is generally unspecialized, with primitive unmolarized P 4 and occasional presence of small lingual cingulum in upper molars [89]. The phylogenetic relationship between Praedama and Megaloceros was accepted by van der Made and Tong [90], however, the cited authors doubted that P. savini could be a direct ancestral form of M. giganteus because of the divergence in mastication adaptations. Vislobokova [23,26,91] regards Praedama as a side phylogenetic branch of the Arvernoceros-Megaloceros lineage. ...
... The distal portion of P. savini is simplified and looks very different from the pattern seen in M. giganteus. The crown part of P. savini antlers is often described as one or two subsequent bifurcations that form a rather simple crown [26,90]. According to Vislobokova [26], the dichotomous pattern of antler crown in P. savini rather reminds one of the antler bauplan of Sinomegaceros. ...
The article presents a preliminary morphological description of the holotype of Mega-loceros giganteus (Blumenbach, 1799) that serves for the description of the species. The article proposes a taxonomical and morphological revision of the nominotypical subspecies M. giganteus gi-ganteus and morphological comparison with other subspecies of M. giganteus. The cluster analysis of diagnostic craniodental and antler characters revealed the systematic position and phylogenetic relationships of M. giganteus with other cervid groups. The genus Praedama is regarded as a closely related phylogenetic branch that linked to the direct cursorial forerunner of Megaloceros that evolved in the middle latitudes of Western Siberia and northern Kazakhstan. The genus Dama has a distant relationship with Megaloceros and represents an earlier phylogenetic branch that evolved in the Ponto-Mediterranean area. The article discusses the secondary adaptations of M. giganteus forms to forest and woodland habitats in Europe and general paleobiogeographic features of the Megaloceros lineage.
... Although Steensma (1988) did not refer these finds to a genus, later authors identified them as Megaloceros aff. savini (Dawkins 1887) (van der Made andTong 2008;Vislobokova 2012). Due to its morphologic peculiarities, this species is often classified in the separate genus Praedama Portis, 1920(e.g., Vislobokova 2012Croitor 2014Croitor , 2018b). ...
... The recently described species Megaloceros novocarthaginiensis van der Made, 2015, has also Praedama morphology (van der Made 2015; Croitor 2018b). Distribution Antler and cranial specimens morphologically similar to P. savini are present in the faunas of Libákos and Kapetánios (Steensma 1988, pp. 168-175;van der Made and Tong 2008). ...
... It is known in the fossil record from the late Middle Pleistocene to the early Holocene, but most of the available samples date from the very last part of its biochronologic range. In the current taxonomic practice, the genus is essentially monotypic (van der Made and Tong 2008;Vislobokova 2012Vislobokova , 2013Croitor 2014), although Croitor (2018b) and van der Made (2019) added two additional species from the Middle East and Spain respectively, while Praedama savini is often included in Megaloceros (e.g., van der Made 2019). Despite its antiquity, the name Megaloceros (emended spelling of the original Megalocerus) has only recently started to be used systematically in the literature, since the recognition of its priority over the previously commonly used Megaceros Owen, 1844 (Lister 1987b; see also Simpson 1945, p. 154). ...
The paleontological research during the last 160 years in Greece has recovered continental deer fossils from at least 100 localities, which span geochronologically from the late Miocene to the end of the Pleistocene, while scanty dental remains date from the middle Miocene. The following species are documented: Procapreolus pentelici (late Miocene), Procapreolus cusanus, Croizetoceros ramosus, Metacervocerus rhenanus, Eucladoceros cf. ctenoides, Rucervus gigans, Praedama aff. savini, Dama vallonnetensis (late Pliocene and Early Pleistocene), Praemegaceros pliotarantoides, Praemegaceros verticornis, Alces latifrons, Megaloceros giganteus, Cervus elaphus, Dama dama, Capreolus capreolus, “Cervus” peloponnesiacus (Middle and Late Pleistocene), while several other samples are incomplete and not identifiable to the species level. Despite their comparatively scarce representation in the fossil faunas, the cervids became more common in time periods with cooler climate, like during the Late Pleistocene.
... The postcranial bones were identified according to the method employed by Brown and Gustafson (1979), Prummel (1988). The specimens were measured according to the methods used by von den Driesch (1976), van der Made and Tong (2008) and van der Made (2015). The linear measurements were taken with sliding caliper in millimeters. ...
... The metapodials of E. boulei are larger than those of the extant species of Cervus, and also larger than those of most co-generic species and even some species of Megacerines; moreover, the metapodials of E. boulei are relatively gracile (van der Made and Tong, 2008;van der Made, 2015, in press). ...
... The pachyostosed mandible in some cervids has been widely noticed (Lister, 1994;Kahlke, 1997;van der Made and Tong, 2008;Vislobokova, 2013;Croitor, 2016), such as Megaloceros, Praemegaceros, and Megaceroides. The Megaloceros species are characterized by the well-known pachyostosis of the skull and, in particular, of the horizontal mandibular ramus. ...
Recent excavations at Shanshenmiaozui site in Nihewan Basin of North China uncovered a rich collection of comb-antlered deer, which includes the first discovery of the yearling antler, complete upper and lower dentitions (both deciduous and permanent), associated limb bones including the longest metapodials ever recovered. Based on toothrow length and the dental characters as well as the postcranial bones, the new fossil materials can be referred to Eucladoceros boulei that is estimated to be 350 kg and represents the largest Pleistocene cervid ever recovered in China. Besides the large size, E. boulei is also characterized by the pronounced anterior cingulids and entostylid ribs on lower molars as well as the moderately pachyostosed mandibles. The present study shows that the body weight of large cervids can be estimated by the length of toothrow and metacarpal, with exception for Cervus elaphus, which has larger toothrow length, but shorter metacarpal and smaller body size. E. boulei is a typical element of the Early Pleistocene fauna in northern China. The early Villafranchian is a bottleneck period for cervid evolution in northern China, which is characterized by the following features: decrease of cervid diversity, disappearance of archaic groups, and the rise of the medium to large-sized three-tined cervini taxa. The sudden appearance of the very large and/or multi-tined cervids at the Pliocene-Pleistocene transition may represent a great migration event of mammals. The Early Pleistocene cervids from Nihewan Basin are very diverse, and are in need of more taxonomic work.
... The postcranial bones were identified according to the method employed by Brown and Gustafson (1979), Prummel (1988). The specimens were measured according to the methods used by von den Driesch (1976), van der Made and Tong (2008) and van der Made (2015). The linear measurements were taken with sliding caliper in millimeters. ...
... The metapodials of E. boulei are larger than those of the extant species of Cervus, and also larger than those of most co-generic species and even some species of Megacerines; moreover, the metapodials of E. boulei are relatively gracile (van der Made and Tong, 2008;van der Made, 2015, in press). ...
... The pachyostosed mandible in some cervids has been widely noticed (Lister, 1994;Kahlke, 1997;van der Made and Tong, 2008;Vislobokova, 2013;Croitor, 2016), such as Megaloceros, Praemegaceros, and Megaceroides. The Megaloceros species are characterized by the well-known pachyostosis of the skull and, in particular, of the horizontal mandibular ramus. ...
Recent excavations at Shanshenmiaozui site in Nihewan Basin of North China uncovered a rich collection of comb-antlered deer, which includes the first discovery of the yearling antler, complete upper and lower dentitions (both deciduous and permanent), associated limb bones including the longest metapodials ever recovered. Based on toothrow length and the dental characters as well as the postcranial bones, the new fossil materials can be referred to Eucladoceros boulei that is estimated to be 350 kg and represents the largest Pleistocene cervid ever recovered in China. Besides the large size, E. boulei is also characterized by the pronounced anterior cingulids and entostylid ribs on lower molars as well as the moderately pachyostosed mandibles. The present study shows that the body weight of large cervids can be estimated by the length of toothrow and metacarpal, with exception for Cervus elaphus, which has larger toothrow length, but shorter metacarpal and smaller body size. E. boulei is a typical element of the Early Pleistocene fauna in northern China. The early Villafranchian is a bottleneck period for cervid evolution in northern China, which is characterized by the following features: decrease of cervid diversity, disappearance of archaic groups, and the rise of the medium to large-sized three-tined cervini taxa. The sudden appearance of the very large and/or multi-tined cervids at the Pliocene–Pleistocene transition may represent a great migration event of mammals. The Early Pleistocene cervids from Nihewan Basin are very diverse, and are in need of more taxonomic work.
... Some fossils of a small species of Megaloceros from the Manzanares valley were described and assigned to Praedama sp. or Dolichodoryceros savini (De Andrés & Aguirre, 1975;Aguirre, 1989), but many more went un noticed and have been classified as red or fallow deer. Remains from up to 10 localities and levels were identified and included in a study on the relationship between the giant deer Megaloceros and Sinomegaceros (Van der Made and Tong, 2008). This study presented a phylogeny of Megaloceros with five species, including M. giganteus, M. savini and three species which were not named. ...
... For the detailed description of dental morphology the nomenclature proposed by Van der Made (1996) is used (Fig. 2). The measurements are given in mm (Tables 1-5 Made and Tong, 2008;radius, astragalus, phalanges: Van der Made, 1996). ...
... Grazers tend to have the condyle in a more elevated position above the occlusal surface, while browsers tend to have it in a lower position. Here a method is used to describe the position of the condyle (Van der Made and Tong, 2008). A reference line is the line through the lowest points of the valleys between the anterior and posterior lobes of the M 1 and M 3 . ...
The Irish elk or Megaloceros giganteus is an emblematic species of the Pleistocene, but its relatives are much less known and were believed to have gone extinct when the Irish elk dispersed into Europe. The species Megaloceros matritensis n. sp. is described here on the basis of material from ten localities and levels in a terrace of the Manzanares river, South of Madrid. It must have been a common species when it lived there some 300–400 ka ago, being contemporary of M. giganteus. This species acquired features, such as enlarged premolars, very thick molar enamel, and a low mandibular condyle, which are masticatory adaptations to an, as yet, unknown diet. The species itself formed part of the diet of people which lived in the area. Megaloceros matritensis fossils are found associated to stone tools of late Acheulean and early Mousterian type. If found in other areas, this species could be indicative for this transitional period. Giant deer might be expected to be good examples of Cope's rule, which holds that species tend to evolve larger body sizes. However, M. matritensis is the last member of a lineage which gradually decreased in size during the Middle Pleistocene. Contrary to what one might expect, size decrease is not un-common in the giant deer.
... This is a poorly understood genus from the final stage of Early Pleistocene and the Middle Pleistocene of Europe. This genus is often regarded as a direct forerunner of the giant deer Megaloceros (Azzaroli 1953;Vislobokova 1990;van der Made & Tong 2008) or its sister form (Lister 1994) and sometimes is synonymized with it (Lister 1987(Lister , 1994van der Made 2013). ...
... Azzaroli (1953) regarded the flattened basal tine of antler as an argument for the phyletic relationship between P. savini and M. giganteus. This assumption was uncritically accepted (Lister 1994;Vislobokova 1990Vislobokova , 2012van der Made & Tong 2008). However, the hypothesized phylogenetic relationship between the two genera based on a single antler character is not safe. ...
... Arvernoceros from Late Pliocene of Europe shows a remarkable bauplan of antlers shared with modern Rucervus and Sinomegaceros (van der Made & Tong 2008;Croitor 2009Croitor , 2016. Unlike Rucervus, Arvernoceros and Sinomegaceros evolved a small distal palmation. ...
... However, a simple scattered diagram of mandible proportions shows that the mandible shape in large-sized Praemegaceros is very similar to the morphological condition found in Eucladoceros and Dama (Croitor, 2006). Van der Made and Tong (2008) found signs of mandibular pachyostosis in a wide variety of cervids and assumed that this specific character evolved among cervids several times in parallel and denied its plesiomorphic significance for the phylogenetic group of giant deer. Therefore, the sporadic occurrence of mandibular pachyostosis in various cervid lineages can not be used as a meaningful taxonomic character at the tribe level. ...
... However, Bärmann and Sanchez-Villagra (2011) report the high number of cranial suture fusion also for some other ruminant genera (Ocapia, Tragelaphus, Kobus, and Antilocapra), seeking the explanation in biomechanical factors. Van der Made and Tong (2008) remarked that the function of temporary storage of minerals should be followed by signs of resorption in pachyostotic mandibles. Actually, even non-pachyostotic bones represent a dynamic system constantly undergoing resorption and deposition of minerals and no particular "scars" of resorption on bone tissue could be seen, taking apart the cases of pathology (Alberts et al., 1983). ...
... At least superficially, the skull shape of Sinomegaceros pachyosteus with its little flexed braincase, broad orbito-frontal part, and short splanchnocranium seems to be a less accentuated version of Megaceroides algericus. Megaceroides is characterized by the presence of a well-developed middle tine, which normally is lacking in Sinomegaceros (Shikama and Okafuji, 1958;Shikama and Tsugawa, 1962;van der Made and Tong 2008). Van der Made and Tong (2008) regard the absence of the middle tine as an essential diagnostic character of Sinomegaceros that is shared with Arvernoceros. ...
The unusual cranial morphology of the endemic extinct African deer Megaceroides algericus (Lydekker,
1890) is described. Some details of cranial and dental morphology suggest that M. algericus is closely
related to the Eurasian giant deer Megaloceros giganteus (Blumenbach, 1799). The paper presents also a
discussion on paleoecology and functional morphology of M. algericus, as well as its origin, phylogenetic
and taxonomic position. Megaloceros mugarensis (Di Stefano, 1996) from the middle Pleistocene of
Levant is regarded as a probable forerunner of M. algericus.
... Ma (Agustí et al., 1987), 1.5-0.7 Ma (Carbonell et al., 1981), Middle Pleistocene (Pons-Moyà & Moyà -Solà, 1979), 0.98-0.79 Ma (Martin, 2012) and around 0.5 Ma (Crégut-Bonnoure, 1999; Van der Made, 2001; Van der Made & Tong, 2008) . Paleomagnetic research suggests, that the fossils come from sediments with reversed polarity, just below the change to normal polarity (Gibert Clols et al., 1999;Ferràndez-Cañadell et al., 2014). ...
... Of the eleven species in the "Megaceros giganteus group", four are currently included in the East Asian genus Sinomegaceros (e.g. Vislobokova, 1990Vislobokova, , 2012R.D. kahlke, 1995Van der Made & Tong, 2008), five were implicitly included in the species Megaloceros giganteus and one in M. savini (Lister, 1993(Lister, , 1994R.D. kahlke, 1995 and one of the species was un-named (and dubious) and has further been ignored. At present, only two named species are included in Megaloceros (e.g. ...
... Delpech & Guérin, 1996;Van der Made, 2005). However, Van der Made & Tong (2008), largely based on biometrics, indicated the existence of three not named species, which are different from M. savini and M. giganteus. One of these was recognized on the basis of material from Cueva Victoria. ...
... The way of measuring the teeth of the Suiformes and the bones of the Artiodactyla is generally after Van der Made (1996). Bovid and cervid metapodials and teeth are measured after Van der Made and Tong (2008) and , 2012. ...
... 3.6.3. Remarks on context and taxonomy Fig. 10 gives the temporal distribution of the giant deer species of the main land of Europe (after Van der Made and Tong, 2008; Van der Made & Dimetrijevi c, in press). Insular species are not considered here. ...
... The first and second phalanges are robust as in Megaceroides and differ in this from Eucladoceros. As far as known, the earlier Megaloceros species had gracile metapodials (Van der Made and Tong, 2008) and first phalanges. The bones and teeth described in this section belong to a large deer, but differ in size from the smaller Eucladoceros species and from Megaloceros spp. ...
An extraordinary sequence of fossiliferous levels at the locality of Gran Dolina in the Atapuerca Hills (Burgos, Spain) records the Early–Middle Pleistocene transition. These levels are well dated by a variety of methods, including palaeomagnetism, which locates Lower–Middle Pleistocene boundary at the top of level TD7. Level TD6 is the type site of the species Homo antecessor and yielded over 90% of the European Early Pleistocene human record, while other levels have an archaeological record. The present paper deals with the earliest Middle Pleistocene ungulates of TD8, but we plan to describe the faunas, or at least the ungulates, of levels TD4 to TD8 in the context of the faunal changes that took place around the Early–Middle Pleistocene transition.
... The latter species is present in Spain during the early Middle Pleistocene (approximately MIS 19 to 14; Atapuerca TD inf.-Arenero Manuel Soto; Made and Tong, 2008); in Germany it is recorded at Voigtstedt and Süssenborn (approximately MIS 17-16) (Kahlke, 1965(Kahlke, , 1969 while in England it is reported at Pakefield, Trimmingham, West Runton and Mundesley (approximately MIS 18 to 15) (Made and Tong, 2008;. A giant deer with palmate brow tine ascribed to M. aff. ...
... The latter species is present in Spain during the early Middle Pleistocene (approximately MIS 19 to 14; Atapuerca TD inf.-Arenero Manuel Soto; Made and Tong, 2008); in Germany it is recorded at Voigtstedt and Süssenborn (approximately MIS 17-16) (Kahlke, 1965(Kahlke, , 1969 while in England it is reported at Pakefield, Trimmingham, West Runton and Mundesley (approximately MIS 18 to 15) (Made and Tong, 2008;. A giant deer with palmate brow tine ascribed to M. aff. ...
... A giant deer with palmate brow tine ascribed to M. aff. savini is recorded at Libakos (Greece; ca 1.2 Ma) and is probably at the origin of M. savini in Western Europe (Made and Tong, 2008). Both P. verticornis and M. savini disappear from the Italian Peninsula before the Isernia FU, about in concomitance with the first occurrence of the genus Dama with distal palmations on the antlers. ...
The Ponte Galeria area is part of the larger “Campagna Romana” and hosted the inner delta of the Tiber River since the beginning of the Middle Pleistocene, allowing the deposition of a series of glacio-eustatically controlled fluvial-lacustrine sedimentary successions very rich in fossil mammal remains. Due to its richness of fossiliferous sites, this area represents an exceptional archive for the study and understanding of the biochronological, paleobiogeographical and paleoenvironmental evolution during the Middle Pleistocene in Europe. A series of recent studies, using the 40Ar/39Ar ages of tephra intercalated within the sedimentary deposits, provided a large amount of geochronological data linking these aggradational successions to the different Marine Isotopic Stages. In the present paper we present a complete review of the faunal assemblages identified so far in the Ponte Galeria area, and we constrain the ages of the faunal units by placing them within the new chronostratigraphic scheme. By means of this interdisciplinary approach, we re-calibrate the mammal assemblages collected in this area and reconstruct a new biochronological and paleobiogeographic framework for the Italian peninsula during the Middle Pleistocene.
... The giant deer (Megaloceros giganteus Blumenbach 1799) was widespread in Pleistocene Eurasia (Stuart et al., 2004;Van der Made, 2006;Pushkina, 2007;Van der Made and Tong, 2008;Shpansky, 2011;Vislobokova, 2011Vislobokova, , 2012. Giant deer antlers are robust, palmate, and extend laterally (Van der Made and Tong, 2008;Stuart et al., 2004), often spanning 3.5 m or more and weighing up to 45 kg (Lister, 1994). ...
... The giant deer (Megaloceros giganteus Blumenbach 1799) was widespread in Pleistocene Eurasia (Stuart et al., 2004;Van der Made, 2006;Pushkina, 2007;Van der Made and Tong, 2008;Shpansky, 2011;Vislobokova, 2011Vislobokova, , 2012. Giant deer antlers are robust, palmate, and extend laterally (Van der Made and Tong, 2008;Stuart et al., 2004), often spanning 3.5 m or more and weighing up to 45 kg (Lister, 1994). The roles of such large antlers have been discussed (Barnosky, 1985;Stuart et al., 2004;Benton and Harper, 2009). ...
... Morphological (Breda, 2005), paleoecological (Van der Made and Tong, 2008;Chritz et al., 2009), paleobiogeographical (Stuart et al., 2004), evolutionary (Vislobokova, 2012), and phylogenetic (Lister et al., 2005;Hughes et al., 2006;Van der Made and Tong, 2008) studies, among others, of giant deer have been published. However, morphological reports have not included the histological detail of normal and pathological antler tissue. ...
We evaluated the skull of an ancient giant deer with a deformity of one antler. The skull was found in the 1930s in the San River near Barycz, in southeastern Poland. Its
dating (39,800±1000 yr BP) corresponds to MIS-3, when the giant deer was widespread
in Europe. Our diagnostics for the antler included gross morphology, radiography,
computed tomography, and histopathology. We noted signs of fracture healing of
the affected antler, including disordered arrangement of lamellae, absence of osteons,
and numerous Volkmann’s canals remaining after blood vessel loss. The antler deformity
appears to be of traumatic origin, with a healing component. No similar evaluation
process has been described previously for this species.
... The giant deer (Megaloceros giganteus Blumenbach 1799) was widespread in Pleistocene Eurasia (Stuart et al., 2004;Van der Made, 2006;Pushkina, 2007;Van der Made and Tong, 2008;Shpansky, 2011;Vislobokova, 2011Vislobokova, , 2012. Giant deer antlers are robust, palmate, and extend laterally (Van der Made and Tong, 2008;Stuart et al., 2004), often spanning 3.5 m or more and weighing up to 45 kg (Lister, 1994). ...
... The giant deer (Megaloceros giganteus Blumenbach 1799) was widespread in Pleistocene Eurasia (Stuart et al., 2004;Van der Made, 2006;Pushkina, 2007;Van der Made and Tong, 2008;Shpansky, 2011;Vislobokova, 2011Vislobokova, , 2012. Giant deer antlers are robust, palmate, and extend laterally (Van der Made and Tong, 2008;Stuart et al., 2004), often spanning 3.5 m or more and weighing up to 45 kg (Lister, 1994). The roles of such large antlers have been discussed (Barnosky, 1985;Stuart et al., 2004;Benton and Harper, 2009). ...
... Morphological (Breda, 2005), paleoecological (Van der Made and Tong, 2008;Chritz et al., 2009), paleobiogeographical (Stuart et al., 2004), evolutionary (Vislobokova, 2012), and phylogenetic (Lister et al., 2005;Hughes et al., 2006;Van der Made and Tong, 2008) studies, among others, of giant deer have been published. However, morphological reports have not included the histological detail of normal and pathological antler tissue. ...
We evaluated the skull of an ancient giant deer with a deformity of one antler. The
skull was found in the 1930s in the San River near Barycz, in southeastern Poland. Its
dating (39,800±1000 yr BP) corresponds to MIS-3, when the giant deer was widespread
in Europe. Our diagnostics for the antler included gross morphology, radiography,
computed tomography, and histopathology. We noted signs of fracture healing of
the affected antler, including disordered arrangement of lamellae, absence of osteons,
and numerous Volkmann’s canals remaining after blood vessel loss. The antler deformity
appears to be of traumatic origin, with a healing component. No similar evaluation
process has been described previously for this species.
... 5644/3367 is similar to the antler of a deer from Libakos (MNQ19) in Greece, which is assigned to Megaloceros sp. (Van der Made and Tong, 2008;Van der Made, 2015) or Praedama sp. (P. ...
... Before these finds, the giant deer from Libakos (MNQ19, ca. 1.3 Ma) from Greece was considered the oldest representative of the genus Megaloceros (s.l.) in Europe (Van der Made and Tong, 2008;Van der Made, 2015. According to J. van der Made, the deer from Libakos might be at the base of the divergence of two lineages: one led to M. savini (=Praedama savini) and the other to M. giganteus (Van der Made, 2015, text-fig. ...
... Both S. tadzhikistanis and S. konwanlinensis are characterized by the rather long beam segment between the basal tine and the distal palmation that may be regarded in this case as an advanced evolutionary character. The genus Sinomegaceros became very successful during the Middle to Late Pleistocene when it was presented by several quite specialized giant species and forms on the vast area from Japan to Central Asia (Shikama and Tsugawa, 1962;Qi, 1983;Ji, 1988;Huang et al., 1989;Sotnikova and Vislobokova, 1990;Vislobokova, 1990Vislobokova, , 2012aVislobokova, , b, 2013Van der Made and Tong, 2008;Okumura et al., 2016). ...
... Interestingly enough, "Dama" sericus from the Late Tertiary of China is nested together with Praesinomegaceros and Sinomegaceros. In contrary to the broadly accepted opinion that Praesinomegaceros, Sinomegaceros, Megaloceros, and Rucervus (Arvernoceros) represent a natural systematic group of Megacerini (Vislobokova, 1990;2012a;b, 2013;Van der Made and Tong, 2008), our multivariate analysis shows that the Praesinomegaceros/Sinomegaceros lineage is morphologically quite distant from Megaloceros and Rucervus (Arvernoceros). The obtained results also indicate that the cranial pachyostosis evolved in Megaloceros and Sinomegaceros represent a physiological evolutionary parallelism (see discussion in Croitor, 2016) and does not have a taxonomic value above the genus level. ...
The article presents a description of a new species of the genus Praesinomegaceros from the Early Pleistocene of Pabbi Hills (Upper Siwaliks) in Pakistan. This is a first record of Praesinomegaceros in the geological past of the Indian Subcontinent. Unlike Praesinomegaceros from the Late Miocene of Central Asia, the Siwalik form is characterized by the loss of basal tine palmation and diminished number of crown ramifications. However, the new Siwalik cervid maintains the characteristic of Praesinomegaceros general antler bauplan. The antler shape of the new species from Pakistan reveals important evolutionary changes in ecology and ethology. Antlers of Praesinomegaceros from Siwaliks have lost their function of social display and “returned” to the effective weapon antler type. The multivariate cluster analysis of antler characters supports the close evolutionary relationship of the new cervid species from the Siwaliks with Praesinomegaceros and Sinomegaceros from the Pleistocene of Asia.
... These specimens were measured with an electronic calliper with 0.01 mm precision. The measuring scheme and abbreviations follow von den Driesch (1976) for horses, deer, and bisons, van der Made and Tong (2008) for the giant deer, as well as Made (2010) for the rhino remains. All values are presented in millimetres. ...
... Family Tables 3 and 4. (in mm, according to the scheme in Made and Tong, 2008) Remarks. Most of the giant deer antlers from Kaniv are characterised by large rosettes, which are among the largest in Eurasia (Fig. S7) (Croitor et al., 2014;Malikov, 2015;Shpansky, 2011Shpansky, , 2014Stefaniak, 2015). ...
Subfossil remains of the Quaternary megafauna from two Ukrainian natural history collections stored at Kaniv Nature Reserve and the National Museum of Natural History NAS of Ukraine were examined. Most of the bones were collected in 1965–1966, in the building pit and the gateway of the Kaniv hydroelectric power plant. The fauna is represented at least by ten taxa (Gulo gulo, Mammuthus trogonterii chosaricus, Mammuthus intermedius, Coelodonta antiquitatis, Megaloceros giganteus, Alces alces, Bison priscus, Cervus elaphus, Capra cf. ibex, and Equus ferus). Skeletal parts are represented mainly by skulls, long bones, horns, and tusks. Based on the species composition of proboscideans, at least part of the faunal assemblage is dated by the end of the Middle Pleistocene (Dnieper Stage = Saale, Warta, MIS 6), but majority of bones could be older or younger (Late Pleistocene and Holocene age) due to the alluvial origin of accumulation.
... Conventional methods were used in the morphological studies, based on visual comparisons and simple morphometrics. The measurements of the Equidae are taken as indicated by Eisenmann et al. (1988), those of the Rhinocerotidae as indicated by Van der Made (2010a), those of the Artiodactyla as indicated by Van der Made (1996) and Van der Made and Tong (2008), and those of the carnivores are taken in a comparable way. All measurements are given in mm, unless indicated otherwise. ...
... All measurements are given in mm, unless indicated otherwise. The measurements are indicated by same abbreviations as used by Van der Made (1996) and Van der Made and Tong (2008). DAP, DT, DMD, DLL mean respectively antero-posterior, transverse, Table 6.1 Faunal list provided by Rivals (2004) based on Aliev's (1969) and Lioubine's (2002) identifications (material hosted in the Medical University of Baku (Azerbaijan) from 1960 to 1989 seasons lead by Huseinov) medio-distal and labio-lingual diametre respectively. ...
During the 1960s to 1980s a human mandible, together with fossils of other animals and a lithic industry, were recovered from Units I to VI of Azokh Cave. After the year 2002, new excavations in Units I to V were undertaken. The new large mammal fossils are described and the fauna is revised, using part of the older collections. The only clear break in the sequence is the appearance of domestic mammals in Unit I. The following taxa recovered from Pleistocenic sediments were identified: Ursus spelaeus (the most abundant), Ursus sp. (U. aff. arctos/thibetanus), Vulpes vulpes, Canis aureus, Canis lupus, Meles meles, Martes cf. foina, Crocuta crocuta, Felis chaus, Panthera pardus, Equus hydruntinus, Equus ferus, Stephanorhinus hemitoechus, Stephanorhinus kirchbergensis, Sus scrofa, Capreolus pygargus, Dama aff. peleponesiaca, Dama sp., Megaloceros solilhacus, Cervus elaphus, Bison schoetensacki, Ovis ammon, Capra aegagrus and Saiga. Most species present are common in western Eurasia. All fossiliferous Units have taxa that in mid-latitude Europe are considered to be “interglacial” elements, while there are no clear “glacial” elements, which suggests temperate conditions despite the altitude of the cave. The evolutionary levels of various species suggest ages of about 300 ka for Units VI–IV, while Units III–II are slightly younger. Domestic mammals indicate a Holocene age for Unit I. Most sediments represent a normal transition between units. Processes of erosion, however, affected the top of the Pleistocene sediments recorded in the cave. Therefore, Unit I (Holocene sediments containing domestic animals) lies disconformably over Unit II (Late Pleistocene).
... Opinions differ concerning the time of origin and early forms of this taxon (Vislobokova, 2012a). Some researchers (Made and Tong, 2008) believe that one of early forms of the genus is Megaloceros aff. savini (Praedama aff. ...
... The antler from the Georgievsk sand pit resembles somewhat those of some members of the genus Sinomegaceros, which also possess a well devel oped palmation. In the palmation development and absence of the middle tine, the deer from Ciscaucasia is most similar to Early Pleistocene Sinomegaceros fla bellatus Teilhard and Middle Pleistocene S. pachyos teus Young (Young, 1932;Teilhard de Chardin, 1936;Vislobokova, 2012) from the Chinese Zhoukoudian locality; but differs from it in the smaller weaker devi ation of the beam, greater development of three poste rior tines, and two flattened and doubled anterior tines of the palmation and also in the more vertical position of the palmation relative to the sagittal skull axis (Made and Tong, 2008;Vislobokova, 2012a). Proba bly, in the antler considered, the first supraorbital tine was probably less massive. ...
A new deer species, Megaloceros stavropolensis sp. nov., from the pre-Apsheronian sandy–clayey deltaic deposits of the Georgievsk sand pit (village of Podgornoe, Stavropol Region) is described. The fauna of large and small mammals from the bone beds of this locality is dated terminal Villanian, end of the MNQ 17 Zone, beginning of the Late Villafranchian, and assigned to the beginning of the Psekups Mammalian Assemblage. This is the earliest known member of the genus, which possesses antlers with well-developed palmation. The middle tine is absent and the posterior tine is a part of the antler palmation. Other specimens of Early Pleistocene Megaloceros are represented by fragments.
... Hadjouis instead considered M. algericus as a possible representative of an older Asiatic form similar to Sinomegaceros pachyosteus recognized in Zhoukoudian (Young, 1932;Dietrich, 1933). Van der Made and Tong (2008) indicated that Sinomegaceros, which is an exclusively Asiatic genus, and its related species, evolved independently without dispersals between east and west Eurasia, and consequently affirmed the absence of a phylogenic relationship with Megaloceros. As seen above, with the different uses of the genus Megaceroides in the literature, the taxonomic and systematic position of this genus has not yet been resolved within the Megacerinae sub-family and a possible ancestor of African origin can be ruled out. ...
... In addition, in Pleistocene lineages of cervids subject to marked seasonality, greater differences in the development of pachyostosis should be expected between males and females, but this has not been yet demonstrated (Lister, 1994) with the exception of M. giganteus. Moreover, we do not yet know whether the annually shed cranial appendages were present in both sexes in the earliest forms of cervids or only in males (Van der Made and Tong, 2008;Davis et al., 2011). Given the current state of knowledge, we cannot reach a clear conclusion concerning the respective relationship between the evolution of the seasonal development of antlers, bone mandibular microstructure, and the physiological processes involved in the development of mandibular pachyostosis of M. algericus. ...
During the course of archaeological test excavations carried out in 2007 in the cave of Bizmoune (Essaouira region, Morocco), seven archaeological layers yielding Pleistocene and Holocene artefacts and faunal remains were identified. In the layers C4, C3 and C2, respectively from the oldest to the most recent, terrestrial Helicidae mollusk shells (Helix aspersa) were dated by 14C. These layers also contained many fragments of eggshell, belonging to Struthio cf. camelus, associated with mammal remains such as Oryctolagus/Lepus, Gazella sp., Sus scrofa, Ammotragus lervia, Alcelaphus buselaphus, Equus sp., Phacochoerus aethiopicus and an undetermined Caprini. Among these remains, an incomplete mandible of Megaceroides algericus Lydekker, 1890 with M1 and M2 was found in layer C3. The 6641 to 6009 cal BP time range attributed to this layer has provided the most recent date known so far for M. algericus. In this study, we review and contextualize the findings of this particular species both in time and space and discuss its systematic position. We describe the morphology of the typical pachyostosic mandibular bone with the teeth and compare the dimensions with existing data. The assumption of the combined development, on the one hand, of the pachyostosic phenomenon and on the other hand, of the body weight fluctuations and growth of antlers for cervids strongly affected by seasonality is not supported. In order to understand the origin and the extinction of M. algericus, we examined the AMS radiocarbon dates available in the literature and calibrated them with RenDateModel software. Comparisons are then made with sea surface temperatures (e.g. GISP2 δ18O), eustasy and related environmental changes throughout the time span of this species. Based on these data a possible migration route by the Strait of Gibraltar connected with with eustatic rises in sea-level rises are discussed. The speciation-extinction processes for M. algericus and their correlations with climatic shifts on a long time-scale in North Africa (e.g. Heinrich events, 8200 cal BP event) are also considered. Finally, this new discovery in Bizmoune cave clearly shows that M. algericus lasted until the very end of the Epipaleolithic, around 6000 cal BP (middle Holocene), whereas this species was formerly not believed to have survived until the early Epipaleolithic (around 8000 cal BP).
... In the case of the artiodactyls, the nomenclature of the dentition follows Bärmann & Rössner (2011) and Made (1996); the latter is also applied to the nomenclature and measurement of suids. For cervids and bovids the measures were taken according to specifications by Made & Tong (2008) and Made (1989Made ( , 2012, except for Ld for the distal phalanx when pertinent (Von Den Driesch 1976). Apart from the specified nomenclature in Table 1, equid phalanges are as 2FIII, with the Arabic number indicating the number of the phalanx and the roman numeral indicating the digit (also for metapodials). ...
The Lower Pleistocene site of Quibas, in Sierra de Quibas (Murcia, Spain) was discovered in 1994 and has since then provided abundant material of typical Epivillafranchian taxa. This biochron belongs to the Early-Middle Pleistocene transition (1.2 – 0.78 Ma), characterised by a change in orbital cyclicity from a 41 kyr cycle to 100 kyr that intensified the climate and culminated in the most important faunal turnover of the Pleistocene regarding large mammals. The Group of Palaeoanthropology of the National Museum of Natural Sciences (CSIC, Spain) and the Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA, Spain) carried out four field seasons from 2015 to 2018. Here we present the large herbivorous mammals recovered from the field, including the first citation of two taxa new to the locality: Stephanorhinus cf. etruscus and Bison cf. voigtstedtensis. We also provide the first description of previously mentioned taxa: Dama cf. vallonnetensis and Sus sp. Together with the remaining herbivores, the faunal community shows a strong European affinity with some regionalism. Compared with other Iberian localities, the site of Quibas stands out for the lack of hominin fossils or any evidence supporting their presence in the area, a peculiar scenario given that the Early-Middle Pleistocene transition broadly speaking sees the arrival of humans into Europe.
... Bear teeth morphology and the scheme of measurements follow Baryshnikov (2007). The remains of even-toed ungulates were identified and measured according to von den Driesch (1976), Van der Made and Tong (2008), Van der Made (2010), and Stefaniak (2015). ...
The Middle Pleistocene was a period of dynamic changes in Europe. During MIS 11, a number of modern mammal taxa appeared and environmental conditions remained warm and favourable for a relatively long time. The Medzhybizh 1 locality of Ukraine dated to this very period comprises alluvial deposits with rich animal remains, which allow not only to reconstruct the fauna composition, but also to highlight the environmental conditions that dominated at this locality. A revision of the fauna of Medzhybizh 1 locality based on remains of all vertebrate groups revealed a taxonomically diverse fish community (16 species of 11 genera) dominated by cyprinids common for lacustrine or riverine assemblages. Amphibians are represented by 11 species, while the number of reptile and bird remains are less significant. Mammals are the most represented group at the locality, including small mammals (30 species), carnivorans (2 species), and ungulates (5 taxa), the latter dominated by C. elaphus. The taxonomic composition of terrestrial groups indicates temperate climate with boreal-type forests and meadows similar to cold steppe, as well as low wet areas and riparian habitats inhabited by amphibians, reptiles, insectivores, beavers, and various voles. The fish assemblage indicates a partially overgrown but well-aerated water body (lake or slow-flowing river) with sandy-silty bottom. Lithic artefacts found at the Medzhybizh 1 locality contribute to a better understanding of relationships between ancient hominins and faunas during the Middle Pleistocene of Eastern Europe.
... 2-I)；1 件带 p2-m1 齿根和 m2、m3 的左侧下颌骨(XZSF.13；图 2-J)。 表 1 嶂山梅氏犀 m3 的测量值与对比Tab.1 Tooth measurements of S. kirchbergensis from Zhangshan and comparison with those of other rhinosSinomegaceros 属[25] 。Sinomegaceros 属最早的化石记录是距今约 1 Ma 的公王岭大角鹿 S. ...
As one of the few reported Pleistocene faunal assemblages from the lower Huaihe
River Region, this taxonomic study sheds new light on the biostratigraphy of the Zhangshan fauna. Zhangshan is a fossil locality in the lower Huaihe River region, located where the modern climatic transition area lies between North and South China. This fossil locality was accidentally found in the 1950’s during the Xinyihe River construction project, when a few poorly preserved fragments were unearthed. Recently, more identifiable specimens were collected during a test excavation and these are the focus of this study. The material originates from a fossil layer around 0.4 m thick, composed of greyish yellow fine sands and rich with calcic concretions.
The following taxa have been identified: Turtles, Alligator cf. sinensis, Palaeoloxodon
huaihoensis, Equus hemionus, Stephanorhinus kirchbergensis, Sus lydekkeri, Sinomegaceros ordosianus, Cervus (Sika) grayi and Bison priscus. The identified species allow for a reassessment of the current record, including specimens of Alligator cf. sinensis that represents the northernmost record of this species during Pleistocene and remains of Bison priscus, which is the most southern recorded example of this taxon. The Palaeoloxodon materials were identified as P. huaihoensis, based on a lamellae number greater than P. namadicus, and of a bigger size than P. naumanni. The mandible of S. ordosianus is the only specimen with a complete cheek teeth series. Study of these Sinomegaceros materials further confirms that mandible thickness, cheek teeth length and antler size are significantly correlated in evolution; at the end of Sinomegaceros lineage, S. ordosianus has the longest cheek teeth row, the slimmest mandible, as well as more slight antler.
Most of the elements of Zhangshan fauna represent extinct species, however, E. hemionus and S. ordosianus were recorded only in Late Pleistocene. Thus we can put the age of Zhangshan fauna at the early Late Pleistocene. The ecological setting of the Zhangshan fauna likely reflects a mixed habitat of swamp, forest and grassland in a relatively warm and humid climate. Zhangshan locality is also the only reported fossil assemblage studied as of yet from the Malingshan mountains, where dozens of Late Pleistocene Paleolithic sites have been discovered through surveys. Although further study is needed of the relationship between these fossils and artifacts from the site, the present research begins to elucidate the environmental background of human evolution in the Malingshan mountains.
... The giant deer Megaloceros giganteus (Vislobokova, 1990, 2011, 2012a, b, Lister, 1994Gliozzi et al., 1997;Sardella et al., 1998;Valli,2003-2004;Made and Tong, 2008;Made et al., 2014) appeared and spread widely during this time period, but became extinct at the end of the Pleistocene in Europe and in the Holocene in Asia. The elk Cervalces latifrons migrated into North America through Beringia and became widespread there, but disappeared at the Pleistocene/Holocene transition both in Asia and North America. ...
This work summarises the state of the art of the studies on the fossil cervids of Poland, from the Miocene through to the present, with an extensive reference to the faunas of the rest of Eurasia and the World. It deals with more than 5500 remains of 16 species from 68 localities. The study intends to give an overview on the taxa present in Poland. The primary focus is on the evolution and stages of dispersal of Cervidae in the frame of the environmental changes that took place across Central Europe. The authors propose to place the bioevents of cervids in the Western European Plio-Pleistocene mammalian biochronological framework, which includes the Ruscinian, Villafranchian, Galerian and Aurelian Mammal Ages. The cervid remains are shown to have great biochronological significance.
... The dental nomenclature and the way of measuring of the Equidae, Rhinocerotidae and Artiodactyla follow Eisenmann et al. (1988), Van der Made (2010a), Van der Made (1996) and Van der Made and Tong (2008). The measurements of the carnivore remains follow largely the same method. ...
Valdavara 3 is a new early late Pleistocene paleontological and archeological cave site in northwestern Iberia. Over 1400 fossils have been collected, representing about 40 species. The fauna is of interglacial aspect and is in accordance with the OSL dates from the fossiliferous layer, which indicate an age of 103–113 ka. The great taxonomical diversity indicates a varied landscape. A small collection of lithic artifacts was found associated with the fossils, demostrating presence of humans and suggesting short non-residential visits to the cave. The fossiliferous site was predominantly formed by natural processes. Many fossil localities have short or biased faunal lists, but the fossil fauna recovered from Valdavara 3 is remarkably diverse and may reflect the fauna which once lived there.
... The species is reported shortly before the Jaramillo palaeomagnetic event (about 1.4-1.3 Ma BP) in Spain (Barranco Léon 5 and Fuente Nueva 3, V5b FC) (Abbazzi, 2010 ), and during the Jaramillo submagnetochron in Italy (Colle Curti, Coltorti et al., 1998 ) and possibly in France (Saint Prest, Guérin et al., 2003; Vislobokova, 2013, but see Bonifay, 1981 Geraads, 1990; Lister, 1993 Made & Tong, 2008). The species is identified roughly at the same time at (Cal Guardiola Layer CGRD2(Terrassa, Catalonia) (Madurell-Malapeira et al., 2015). ...
This research aims to investigate the large mammal faunal dynamics in SW Europe during the late Early Pleistocene. At that time, the climate forcing known as Mid-Pleistocene Revolution (MPR) induced deep, more or less gradual alterations and latitudinal displacements in European terrestrial biomes and exerted great influence on dispersal and dispersion of mammalian species. Large mammals did not generally move in multi-species waves of dispersal, rather each species changed its range depending on the suitability of environmental conditions in respect to its own environmental tolerances and ecological flexibility. Factors driving the remodelling of the range of a taxon, and time and mode of its dispersal and diffusion into SW Europe differed from species to species as from one territory to another, leading to diachronicity/asynchronicity in local first appearances/lowest local stratigraphical occurrences. As a result, correlations and biochronological assessments of local faunal assemblages may be difficult especially when firm chronological constraints are unavailable. Whether the peculiar composition of the late Early Pleistocene fauna, characterised by the persistence of some Villafranchian species and discrete new appearances of some taxa that will persiste during the Middle Pleistocene, may be indicative of any high rank biochronological unit is discussed. Evidence from SW Europe suggests that the chronological range of the so-called Epivillafranchian biochronological unit (whatever its biochronological rank could be) may span from about 1.5 (i.e. Lowest local Stratigraphical Occurrence, LlSO datum, of, among others, Homo, Xenocyon lycaonoides, Canis ex gr. C. mosbachensis, Megantereon whitei, advanced stenonoid horses, Praemegaceros, Bison) and about 0.85 Ma (i.e. Highest local Stratigraphical Occurrence, HlSO datum, of X. lycaonoides, M. whitei, P. pardoides). The biochronological unit would tentatively correspond from a biostratigraphical point of view to a virtual "range-zone", i.e. the hypothetical body of strata representing the known stratigraphical occurrence in W Europe of X. lycaonoides-M. whitei. Nonetheless, several factors (e.g. the confusing taxonomic treatment of some taxa, the heterogeneous consistency of the fossil record in space and time-particularly as regard the number and richness of local faunal assemblages-, taphonomical biases, and the apparent dyachronicity asynchronicity of LlSO of some species that dispersed toward Western Europe during the post-Olduvai Early Villafranchian) make it difficult any attempt to "formalise" the "Epivillafranchian biochronological unit" as a European Land Mammal Age (ELMA). All in all, several lines of reasoning suggest an informal use of the term Epivillafranchian, pending a complete revision of the Villafranchian ELMA, its subdivisions and of the rank that each subdivision may actually have.
... from the late Early Pleistocene (ca. 1 Ma) site of Cueva Victoria near Cartagena in Murcia (van der Made, 2015a); this species is larger than M. savini which it resembles (cf. van der Made & Tong, 2008 ). Dama cf. ...
Cueva Negra del Estrecho del Río Quípar overlooking the Río Quípar, a Río Segura tributary, is an upland rock-shelter 75 km north of the Mediterranean coast and 110 km west of the Segura river-mouth. It contains undisturbed sediment 5 m deep assigned by magnetostratigraphy to >0.78 Ma (Matuyama magnetochron). Optically stimulated sediment luminescence dating implies ≥0.5 Ma and mammalian biochronology (notably, of Arvicolid rodents) indicates >0.7-<1 Ma. Remains include an "Acheulian" limestone "hand-Axe," and small chert, limestone or quartzite artifacts, knapped on site, often by bipolar reduction or repetitive centripetal flaking of small discoidal cores. Secondarily-flaked ("retouched") artifacts include small irregular chert fragments, resembling chert at an adjacent conglomerate outcrop according to laser-Ablation inductively-coupled plasma mass-spectrometry of 19 lanthanide elements, though some chert may have been obtained from up to 30 km away. Faunal remains and pollen are compatible with mild (plausibly MIS-21), damp, fluvio-lacustrine environments. Evidence of fire in a deep, sealed layer includes thermally-Altered, lustreless and shattered chert, and both charred and white calcined fragments of bone. Taphonomical analysis and electron microscopy of bone fragments attribute discolouration to burning, not to post-depositional mineral staining. Sediment geochemistry and thin-section micromorphology have been undertaken. FTIRS, TL and ESR analysis of chert and bone imply firing temperatures of about 500C at least. Drawing on findings from Cueva Negra, the purpose of this paper is to offer an interpretation of Palaeolithic activity from the perspective of hominin cognitive versatility, techno-manual dexterity, and palaeoeconomic extractive behaviour in long-vanished Western European palaeoecological and palaeobiogeographical contexts.
... Azzaroli (1953) regarded the flattened basal tine of antler as an argument for the phyletic relationship between Praedama savini and Megaloceros giganteus. This point of view was uncritically accepted by other authors (Lister 1987(Lister , 1994Vislobokova 1990Vislobokova , 2012van der Made & Tong 2008). However, the position of basal the beam (the portion between first and middle tines) is somewhat longer than the second segment (the beam portion between the middle tine and the distal bifurcation). ...
This article proposes a brief overview of opinions on cervid systematics and phylogeny, as well as some unresolved taxonomical issues, morphology and systematics of the most important or little known mainland cervid genera and species from Late Miocene and Plio-Pleistocene of Western Eurasia and from Late Pleistocene and Holocene of North Africa. The Late Miocene genera Cervavitus and Pliocervus from Western Eurasia are included in the subfamily Capreolinae. A cervid close to Cervavitus could be a direct forerunner
of the modern genus Alces. The matching of results of molecular phylogeny and data from cervid paleontological record revealed the paleozoogeographical context of origin of modern cervid subfamilies. Subfamilies Capreolinae and Cervinae are regarded as two Late Miocene adaptive radiations within the Palearctic zoogeographic province and Eastern part of Oriental province respectively. The modern
clade of Eurasian Capreolinae is significantly depleted due to climate shifts that repeatedly changed climate-geographic conditions of Northern Eurasia. The clade of Cervinae that evolved in stable subtropical conditions gave several later radiations (including the latest one with Cervus, Rusa, Panolia, and Hyelaphus) and remains generally intact until present days. During Plio-Pleistocene, cervines repeatedly dispersed in Palearctic part of Eurasia, however many of those lineages have become extinct.
This chapter focuses on the evolutionary and paleoecological context of Late Pleistocene large mammal faunas from East Asia. Through the Quaternary, the faunal physiognomy of East Asia has largely set the pretext for the present-day division between the Palearctic and Oriental zoogeographic realms; although recurrent climatic oscillations created faunal distribution patterns that were manifestly counterintuitive with respect to the traditionally understood geographic and environmental envelopes of many mammal taxa. The fossil record now convincingly demonstrates that keystone megafaunal herbivores of the mammoth steppe biome which characterized Middle-Late Pleistocene Northern Eurasia, such as Mammuthus, Coelodonta, and Bison, oversaw important hallmarks of their early adaptive evolution in northern China and the Qinghai-Tibetan Plateau. It is then paradoxical that the mammoth steppe biome only partially extended into northern China and Japan at its maximum expansion during the Last Glacial. Instead, mid-latitude Eastern Palearctic was mainly dominated by a Palaeoloxodon-Coelodonta association also frequented by Sinomegaceros and Stephanorhinus, suggesting closer ecological analogy with the Western Palearctic interglacial fauna demarcated by Palaeoloxodon and Stephanorhinus. In southern China, the Ailuropoda-Stegodon Faunal Complex constituted a mixture of Chinese endemics and species typical of today's Indo-Malayan region (e.g., Pongo and Dicerorhinus), adapting to habitats which became more seasonal, with greater resource variability throughout the Pleistocene.
Massive human-induced declines of large-sized animals and trees (megabiota) from the Late Pleistocene to the Anthropocene have resulted in downsized ecosystems across the globe, in which components and functions have been greatly simplified. In response, active restoration projects of extant large-sized species or functional substitutes are needed at large scales to promote ecological processes that are important for ecosystem self-regulation and biodiversity maintenance. Despite the desired global scope of such projects, they have received little attention in East Asia. Here, we synthesise the biogeographical and ecological knowledge of megabiota in ancient and modern China, with relevant data mostly located in eastern monsoonal China (EMC), aiming to assess its potential for restoring functionally intact ecosystems modulated by megabiota. We found that during the Late Pleistocene, 12 mammalian megafaunal (carnivores ≥15 kg and herbivores ≥500 kg) species disappeared from EMC: one carnivore Crocuta ultima (East Asian spotted hyena) and 11 herbivores including six megaherbivores (≥1000 kg). The relative importance of climate change and humans in driving these losses remains debated, despite accumulating evidence in favour of the latter. Later massive depletion of megafauna and large-sized (45-500 kg) herbivores has been closely associated with agricultural expansion and societal development, especially during the late Holocene. While forests rich in large timber trees (33 taxa in written records) were common in the region 2000-3000 years ago, millennial-long logging has resulted in considerable range contractions and at least 39 threatened species. The wide distribution of C. ultima, which likely favoured open or semi-open habitats (like extant spotted hyenas), suggests the existence of mosaic open and closed vegetation in the Late Pleistocene across EMC, in line with a few pollen-based vegetation reconstructions and potentially, or at least partially, reflecting herbivory by herbivorous megafauna. The widespread loss of megaherbivores may have strongly compromised seed dispersal for both megafruit (fleshy fruits with widths ≥40 mm) and non-megafruit plant species in EMC, especially in terms of extra-long-distance (>10 km) dispersal, which is critical for plant species that rely on effective biotic agents to track rapid climate change. The former occurrence of large mammals and trees have translated into rich material and non-material heritages passed down across generations. Several reintroduction projects have been implemented or are under consideration, with the case of Elaphurus davidianus a notable success in recovering wild populations in the middle reaches of the Yangtze River, although trophic interactions with native carnivorous megafauna have not yet been restored. Lessons of dealing with human-wildlife conflicts are key to public support for maintaining landscapes shared with megafauna and large herbivores in the human-dominated Anthropocene. Meanwhile, potential human-wildlife conflicts, e.g. public health risks, need to be scientifically informed and effectively reduced. The Chinese government's strong commitment to improved policies of ecological protection and restoration (e.g. ecological redlines and national parks) provides a solid foundation for a scaling-up contribution to the global scope needed for solving the crisis of biotic downsizing and ecosystem degradation.
The swollen mandible is a common feature in megacerines, especially in Sinomegaceros pachyosteus, which is likely related to their feeding strategy. Here, we compare the hemimandibular mechanical performances of four megaherbivores, i.e., S. pachyosteus, Alces acles, Bubalus bubalis, and Equus caballus using finite element analysis. The geometric models of the cranium and the hemimandible were reconstructed and the occlusal process was simulated. Full-sized models were analysed for the von Mises stress comparison and volume-scaled models were used for direct comparisons of the strain energy. Both strain energy and maximum von Mises stress of S. pachyosteus are as low as B. bubalis and E. caballus, and significantly lower than those of A. alces. We conclude that S. pachyosteus has the potential to be a grazer, similar to the diet of B. bubalis and E. caballus.
The Chinese open-air site Xuchang-Lingjing (Henan) is located near the border between the Palearctic and the Oriental biozone in a lowland depression at the southern edge of the North China Plain. The site yielded a Middle Palaeolithic assemblage that includes the two fragmented, incomplete human (possibly Denisovan) skulls (Xuchang 1 and Xuchang 2), more than 15,000 artefacts and more than 40,000 mammalian remains representing 20 taxa. The composition of the faunal assemblage is biased by hominin hunting activities; it is, however, diverse and “natural” enough to be used as a proxy to reconstruct the paleoenvironmental condition during the human occupation of the site. The Palearctic faunal assemblage indicates a grassland-dominated palaeoecological environment, with a mosaic of scattered forest and mixed forest vegetation as well as along rivers and/or lakes the occurrence of swampy areas and with bushes. The OSL-dates and the assumed correlation with the last interglacial (MIS 5) palaeosol strongly suggest that we are dealing with an interglacial fauna. However, the fact that the Lingjing assemblage represents the southernmost Palearctic fauna questions the assumption that the Lingjing Middle Palaeolithic finds date from an interglacial period. A late Middle Pleistocene (MIS 6) or Late Pleistocene glacial or stadial phase (MIS 4) seems more likely.
The intra-montane Guadix-Baza Basin is one of the few continental basins in Europe that hosts a well-dated set of fossiliferous sites spanning from the latest Miocene to the late Middle Pleistocene. The Cúllar de Baza 1 (CB-1) represents a key site to investigate the effects of the Early-Middle Pleistocene Transition, considered a fundamental transformation in the Earth’s climate state. Our review and update of the large mammal assemblage, and particularly equids, is of paramount relevance to understand the systematic affinities and the evolution of the Early and Middle Pleistocene European horses. We confirm the occurrence of two different taxa, the medium sized Equus altidens and the larger E. suessenbornensis. Moreover, we illustrate that CB-1 is essential for the biochronological studies of the latest Early Pleistocene/Middle Pleistocene transition (Epivillafranchian/Galerian ELMA); in particular with regard to the Last Occurrences of the Etruscan rhino Stephanorhinus etruscus and the large deer Megaloceros savini and the First Occurrence of the water-rat Arvicola mosbachensis in the Iberian peninsula. Finally, a development of a mosaic environment characterised the CB-1 site contrasting with the conditions reported for other Iberian late Early and Middle Pleistocene localities.
Prehistoric and recent extinctions of large-bodied terrestrial herbivores had significant and lasting impacts on Earth’s ecosystems due to the loss of their distinct trait combinations. The world’s surviving large-bodied avian and mammalian herbivores remain among the most threatened taxa. As such, a greater understanding of the ecological impacts of large herbivore losses is increasingly important. However, comprehensive and ecologically-relevant trait datasets for extinct and extant herbivores are lacking. Here, we present HerbiTraits , a comprehensive functional trait dataset for all late Quaternary terrestrial avian and mammalian herbivores ≥10 kg (545 species). HerbiTraits includes key traits that influence how herbivores interact with ecosystems, namely body mass, diet, fermentation type, habitat use, and limb morphology. Trait data were compiled from 557 sources and comprise the best available knowledge on late Quaternary large-bodied herbivores. HerbiTraits provides a tool for the analysis of herbivore functional diversity both past and present and its effects on Earth’s ecosystems.
Systematic excavation and multidisciplinary research undertaken over three decades have deepened our understanding of the early Palaeolithic archaeology at Cueva Negra del Estrecho del Río Quípar (Caravaca de la Cruz, Murcia, Spain). New results from biochronology and combined ESR and U-series dating corroborate previous magnetostratigraphy, placing the entire excavated sequence between the Jaramillo sub-chron and the Matuyama-Brunhes boundary (i.e. ca. 990-772 thousand years ago (ka)); palaeontological and palynological findings reflect temperate environmental conditions. A bifacially flaked limestone hand axe was excavated 1 m below the top of the Pleistocene sequence. The Equus cf. altidens tooth that provided the ESR estimate was excavated 1 m below the hand axe. Throughout its 5-m-deep sedimentary sequence, small nodules, fragments and struck flakes make up the bulk of the Palaeolithic assemblage. Stratigraphical analysis points to undisturbed continuous sedimentary deposition above a layer of ashy sediment, encountered 4.5 m below the top of the Pleistocene sequence, which contained thermally altered bone and heat-shattered chert cores and flakes. Cueva Negra is among the earliest European sites with firm evidence of combustion.
The destroyed site(s) of Koskobilo (Olazti, Navarre, Northern Iberian Peninsula) have yielded unique archaeopaleontological evidence in the Western Pyrenees region. The quarry uncovered a karstic site with faunal remains in 1940, and fossils were recovered both in situ and from the quarry dump. Ten years later, while the quarry was still working, a new visit to the dump yielded a large lithic assemblage and additional fossil remains
with a different taphonomic pattern, which has been interpreted as the remains coming from a different site or zone within the same karst system. Here we re-study the paleontological evidence and provide new dating on a speleothem covering a Stephanorhinus hemitoechus tooth, which has yielded a minimum date of c. 220 ka for part of the assemblage. In total, the fossil assemblage comprises 38 mammal and six avian taxa and three fish remains. The faunal evidence indicates that in 1940 a mix of taxa from both the Middle and Upper Pleistocene were recovered, and it is difficult to assign most of them to a concrete period. However, based on biochronological criteria some of the identified taxa (e.g., Ursus thibetanus, Ursus cf. deningeri, Cuon cf. priscus, Macaca sylvanus, cf. Megaceroides) could be roughly contemporaneous with the dated rhino tooth, which would provide a new window to the Middle Pleistocene of the region, with deposits from MIS 7d and/or older. Despite thedifficulties in studying this collection, recovered without stratigraphic context and in a salvage operation, Koskobilo has yielded an important paleontological assemblage which helps to understand the paleoecology of the Middle Pleistocene human occupations in the Western Pyrenees.
The article presents a description of cervid remains from the Middle Pleistocene Acheulean site of Gruta da Aroeira (= Galerias Pesadas) in central Portugal. The assemblage comprises the remains of four deer species: Cervus elaphus, Praedama cf. savini, Haploidoceros mediterraneus, and Dama cf. vallonnetensis, making Gruta da Aroeira the first site in the Iberian Peninsula at which the genus Haploidoceros has been documented in the Middle Pleistocene. Virtually all the cervids documented at the site demonstrate a degree of endemism, including evolutionary modifications of skull, antlers and dentition or a reduction in body size. The unusual richness of the cervid community at Gruta da Aroeira may be related to the specific biogeographic conditions of the Middle Pleistocene in Iberia. Thus, while the Iberian Peninsula’s geographical link with the temperate west Eurasian zone facilitated the dispersal of cervids of palearctic origin into Iberia, it impeded the dispersal of ruminants from warmer, more arid areas. The endemic character of the Middle Pleistocene cervids and the biodiversity of the Iberian Peninsula should shed some light on the paleobiogeography of Iberian hominins and their role in hominin hunting or their economic strategies.
Characteristics of the postcranial skeleton of Arvernoceros ardei and Cervus perrieri from Perrier (France, late Pliocene, 2.78 ± 0.01 Ma BP, MNQ 16b) are described and distinguished. This was made possible by using the skeletal remains of A. ardei from Kaltensundheim (Germany, Thuringia, late Pliocene age) as identification keys. C. perrieri did not occur in Kaltensundheim. Twenty-five skeletal differences of A. ardei and C. perrieri are detected, and are compared to the genera Eucladoceros, Cervus, Praemegaceros and Megaloceros. The results demonstrate a high level of conformity of A. ardei with Eucladoceros in predominantly plesiomorphic features; major differences exist between A. ardei and Cervus, and A. ardei and Megaloceros. C. perrieri shows several similarities with Cervus elaphus, in which both differ from A. ardei and Eucladoceros. An association of C. perrieri to the genus Cervus is indicated by the postcranial skeleton, as has been discussed in the literature based on antler characteristics. By the findings of Kaltensundheim, a precise association of dental and skeletal characteristics to A. ardei is possible now. The allocation of E. giulii from Untermaßfeld (Germany, Thuringia, Jaramillo event) to the genus Arvernoceros appears doubtful based on the dental and skeletal features of A. ardei from Kaltensundheim and Perrier. The original classification within the genus Eucladoceros is considered probable.
Extensive fieldwork and detailed studies during the last three decades have enriched our understanding of the Plio-Pleistocene large mammal record of Greece. While the unearthed material is abundant, it is not evenly distributed throughout the Plio-Pleistocene; therefore, there are time intervals in this period for which the known large mammal fauna is limited and our knowledge is poor. The Greek Plio-Pleistocene large mammal record reveals a paleoenvironmental transition from open woodlands in late Pliocene, to savannah-like landscape during the early Pleistocene, and to open grasslands during the late early Pleistocene. During this environmental shift, several taxa arrived in Greece in their westward expansion, whereas others made their last European appearance. The arrival of Homo in Europe is discussed in relation to the Greek faunal record. The available data cannot clearly distinguish between an African or an Asian origin, but the latter is supported by more evidence.
The rich Quaternary fossil record from the Cantabrian region (NW Iberia) is virtually restricted to the Late Pleistocene, with Middle Pleistocene findings very rare. This work presents the study of two Middle Pleistocene palaeontological sites in the Cantabrian region: Llantrales quarry and Mestas de Con mine, both discovered and first described in the 1950s but never studied in detail thereafter. The material was here re-analysed and re-evaluated. Taxonomic attributions have been changed in most cases and, consequently, chronological assignments of the fossils have also been changed. This first detailed study of the assemblage from Llantrales quarry revealed the occurrence of the large-sized deer Praemegaceros solilhacus, which has never been reported previously in North Iberia. Cervus cf. elaphus and Stephanorhinus cf. hundsheimensis were also identified. The age of these fossils was originally regarded as Pontian (late Miocene), but new palaeontological results indicate a much younger age, most likely from around 0.8 to 0.5 Ma. A taxonomic re-evaluation of the fossils from Mestas de Con provided different results with respect to the 1950s works. The faunal association is composed of a large-sized deer (probably corresponding to Praemegaceros), Cervus elaphus, Capreolus cf. capreolus, Bison sp. (small sized), Equus sp. (large sized), Stephanorhinus hemitoechus, Ursus sp. and Homotherim latidens. This faunal association was evaluated within a western European context and proves to be highly analogous to other contemporary assemblages there. The chronology was initially regarded as late Villafranchian (in a broader sense). New results, documenting the co-occurrence of Stephanorhinus hemitoechus and Homotherium latidens, indicate a more precise and younger age, most likely between 0.6 and 0.4 Ma. These two fossil assemblages provide new and relevant information on the Middle Pleistocene faunas from a geographical region where this information, compared with other western European areas, is rare and poorly known.
This research aims to deconstruct time and mode of dispersal of large mammals in SW Europe during the post-Olduvai Early Pleistocene, focusing on asynchronous versus diachronous appearance bioevents in the Iberian Peninsula, France and Italy. The geography of the region and its rich Quaternary fossil record is particularly suitable for studying the role (if any) of geographical/ecological barriers in either preventing some taxa to enlarge their range or causing their delayed appearances in some territories. The database consists of taxonomically revised lists of large mammalian species from selected faunal assemblages (LFAs) ranging from about 1.6 to 0.7 Ma. Results obtained show that a few diachronous appearances permit following the displacement/enlargement of species range limits across the studied region. Conversely it is a changing issue understanding which among several factors (e.g. the different impact of global climate changes in different geographic settings, differences in species resilience to habitat disturbance variability in competition and predation patterns within local faunal complexes, and heterogeneity of the fossil record) might have caused asynchronous appearances. The complex interplay of niche differentiation and biotic interactions in each region may be among the factors causing asynchronicity, though the influence of the heterogeneous consistency of the fossil record in space and time (e.g. number and richness of LFAs included in each FC), and taphonomical biases affecting the chronological reliability of local first/last appearances, cannot be discounted especially as regard to rare species.
Cueva Victoria is a karst-infilling site with early Pleistocene vertebrate fossil remains. It was discovered in the first half of the XXth century by mining explotation. The fossil record of Cueva Victoria is exceptional because of its large biodiversity and because it is the only locality in Europe where the African primate Theropithecus oswaldi occurs. Here, we summarize the history of paleontological research and excavation campaigns. We provide a catalogue of the different locations in the site, and a reference list of the publications on this site.
The fossil remains of cervids from Trlica in Montenegro are described and assigned to the elk or moose Alces cf. carnutorum, the roe deer ?Capreolus sp., the red deer Cervus elaphus, and Eucladoceros. The new species Eucladoceros montenegrensis n. sp. is named on the basis of 169 bones and teeth. The Early and early Middle Pleistocene was a time with a great diversity in giant deer. Their features are discussed, where possible described by morphometrics, and compared to the material from Trlica. On this basis the new species Eucladoceros montenegrensis is named. At present, the oldest well dated record of a red deer (Cervus elaphus) in Europe is from 936 to 990 ka. Various ages have been proposed for Trlica, both younger and older than the Jaramillo Subchron. The age of the locality is discussed.
Remains of the large-sized deer Arvernoceros from the sites of Madonna della Strada and Selvella (late Early Pleistocene, central Italy) are described. At present, these records represent the first occurrence of this genus in Italy.
These remains show morphological and morphometric characters comparable to the species Arvernoceros giulii and quite different from other Early Pleistocene large-sized deer. A. giulii was a typical species of a savannah-like environment and it seems to have evolved on the plains of central Asia. The dispersal of the species into Western Eurasia is probably caused by transformations of the eastern Paratethys during the Early Pleistocene and its occurrence in Italy may also have been favoured by climatic changes and by the geographical position of the peninsula.
In this paper we present a reconstruction of the stratigraphic setting of the continental sedimentary sequences that were deposited by the Paleo-Tiber River within the greater area of Rome between 0.9 and 0.6 Ma, carried out through analyses of a large number of borehole data. Through palinspastic restoration of several cross sections we depict the original geometry of the sedimentary record that has been dislocated by intense tectonic activity linked to volcanism, and we reconstruct the geologic and paleogeographic evolution of this area.
Moreover, we provide a complete review of the chronostratigraphic and magnetostratigraphic data reported in previous work, and we extend paleomagnetic analyses to three new clay sections. These geochronological constraints allow us to compare aggradation of the Paleo-Tiber sedimentary successions with the d18O record, evidencing a strict link between sedimentation and sea-level changes in the Rome area. By doing so, we provide a direct test on the timing of the sea-level rise for MIS 19 through MIS 15: a record of data for which no equivalent exists in the literature.
In the Upper Rhine area, new finds of Dama dama of the last interglacial period belonging to collections of the Naturkundemuseum Stuttgart and the private collections of F. MENGER, H. HARTNAGEL, H. BUCHHAUPT are described. The material is compared to the recent European fallow deer, the Eemian Dama from Lehringen (Niedersachsen, Verden a. d. Aller) and D. dama from Neumark-Nord at the northern periphery of the Geiseltal (Sachsen-Anhalt, SE of Halle a. 4. Saale), which are either of Eemian or intra-Saalian interglacial age. Dama dama from the Upper Rhine area is characterised by a 10% greater body size compared to the recent fallow deer, and a 10% smaller body size in relation to D. dama from Neumark-Nord (Tab. 1, 2, Figs. 14-20). The antlers can be distinguished from D. dama dama and D. dama from Neumark-Nord by strongly developed additional tines above the browtme (Figs. 7-13). D. dama dama and the fallow deer from the Upper Rhine area share short browtines. The browtines of D. dama from Neumark-Nord are much longer (Figs. 1-2). Like in D. dama from Neumark-Nord, the beam can be flattened above the browtine forming a dorsal ridge. The antlers of the fallow deer from the Upper Rhine area morphologically interlink D. dama from Neumark-Nordand the recent European fallow deer.
Lines of arrested growth (LAGs) are a typical feature of the bone histology of ectothermic tetrapods but have received little study in mammals and birds. However, LAGs have figured prominently in the debate about dinosaur physiology. Here we describe the bone histology, including the occurrence of LAGs, in an extensive sample of herbivorous mammals from the Late Pleistocene of Germany, mainly from the Rhine-Herne ship channel. Taxa sampled include the cervids Megaloceros giganteus, Cervus elaphus, and Rangifer tarandus, the bovids Bos primigenius and Bison priscus, the equid Equus sp., the rhinocerotid Coelodonta antiquitatis, and the elephantid Mammuthus primigenius. Samples were preselected for macroscopic evidence of cyclical growth. Bones sampled were mainly metatarsals as well as tibiae and indeterminate long bone fragments. All samples show fibro-lamellar bone in the cortex that is replaced by secondary bone to varying degrees. Most samples show one or more regularly spaced LAGs, sometimes even preserved in secondary bone. Surprisingly, there are distinct differences in the histology of the various taxa in terms of the arrangement of the primary vascular network and the patterns of remodeling. The common development of LAGs in these endothermic Late Pleistocene mammals calls into question the argument that LAGs in dinosaur bone indicate an ectothermic physiology.
The morphology of the lower fourth premolar has been used as a taxonomic indicator in the pecoran fossil record, for example to separate cervid species, and to differentiate giraffids from cervoids. We demonstrate considerable inter- and intraspecific variation in cervoid and bovid P 4morphology, and show in particular that the typical giraffic type of P 4, with suppression of the central connection between labial and lingual walls, appears among living and fossil cervoids, and very exceptionally among bovids. This character of the premolar dentition has been used to unite the lower Miocene pecoran genus Triceromeryx, known from a single incomplete individual from Spain, with the Giraffidae, although it was originally united with the Dromomerycidae on the basis of the possession of an occipital horn. We conclude that the known material of Triceromeryx is insufficient to assign this genus firmly to any known pecoran family, and that it should remain for the time being as Pecora incertae sedis.-Authors
The systematic position of Dama among plesiometacarpal cervids of the Pleistocene and Holocene was studied based on morphological characters of the postcranial skeleton, the teeth and the antlers. The metric analysis provides an overview of skeletal dimensions and functional adaptations in the appendicular skeleton. Contrary to the antler and tooth characters the postcranial characters are suitable for manual as well as computer-aided phylogeny reconstruction. The genus Dama Frisch 1775 can already be distinguished from Cervus L. at the Pliocene-Pleistocene boundary, Dama rhenana (= Cervus rhenanus Dubois 1904) and Dama nestii Azzaroli 1947 are ancestors of Middle Pleistocene and modern fallow deer. In Europe the genus Cervus appears first in the late Villafranchian of Central Italy with Cervus sp. from Capena. The synchronous occurence of Axis sp. in Pirro-Nord documents the immigration of 'new' cervids in the latest Early Pleistocene in Central Italy which replace Dama nestii. The systematic position of Axis in relation to Dama and Cervus remains an open question.
On June 24th, 1797 some magacerid bones were found at a hillside of Fukamachi, Kamikuboiwa, Tomioka City, Gunma Prefecture, and on July 7th of the same year natives excavate the site and gained antlers, jaws, vertebrate, scapula, limb bones etc.; they presented them to MAEDA, the feudal lord of the district. In November, 1798, a small stone monument of this discovery was built upon the hill. In the summer of 1800, Motohiro TANBA, head doctor of the TOKUGAWA family, kept a record of his description, illustration and determination of the materials. His manuscript is very accurate and now preserved in Hensyo temple in Kamikuroiwa. Choyokan Manpitsu written in 1811 by Tozan KAMAHARA mentioned this discovery. The bones had been stored in the house of MAEDA in Edo, but in 1933 Toshisada MAEDA donated all the materials to the Zyagu shrine in Nanukaichi, Tomioka City, and henceforward they have been kept perfectly as sacred treasures of the shrine. In April, 1959 Syuichi TSUGAWA visited the shrine and recognized the treasures as megacerid fossils of excellent preservation. In March, 1961, the writers visited the locality and verified the existence of the stone monument. The materials of this interesting discovery and history is the best specimens ever known in Japan of Sinomegaceroides yabei (Shikama) of late Pleistocene.
The giant deer Megaloceros giganteus is one of the emblematic Pleistocene mammals. Material from various Dutch localities and large samples from Germany and the North Sea were studied and compared with data and results of studies by Lister (1994) and Stuart et al. (2004) and a hypothesis on the evolution and biogeography of the species is proposed. The earliest representatives of this species had relatively long and slender limb bones. During the Eemian, a sudden and important increase in robusticity occurred in the west European populations, while in the east European populations, robusticity increased only a little. The robust branch became extinct around 20 ka ago (uncalibrated radio carbon years). Between 10 to 13 ka, western Europe was colonized from eastern Europe by Megaloceros with limb bones of an intermediate robusticity. These changes in the robusticity of the limbbones are mainly caused by changes in length of the bones and seem to be related to locomotion, while body weight seems to have changed little. © E. Schweizerbart'sche Verlagsbuchhandlung (Nägele u. Obermiller), 2006.
The Ungulates from Atapuerca: Stratigraphy and Biogeography. The Sierra de Atapuerca, near Burgos (Spain), has various fissure fillings that yielded fossil animals, including fossil man, and archaeological remains, of late Early Pleistocene to Holocene age.Level TD6 in the locality Gran Dolina, which contained the type material of Homo antecessor as well as archaeological objects, and levels TDW4 and TDE5 yielded a late Early Pleistocene fauna, including the ungulates: Equus cf. altidens, Stephanorhinus etruscus, Dama ‘nestii’ vallonnetensis, Cervus elaphus, Eucladoceros giulii, cf. Bison voigtstedtensis, Sus scrofa and Mammuthus sp. The lower part of TD8 yielded a fauna that is very similar, and that includes also Hippopotamus. TD7 yielded remains of Ovibos or Praevibos. The lower part of the Sima del Elefante sequence yielded some remains that might represent a fauna similar to that of TD4–6.Levels TD10–11 of Gran Dolina yielded: Equus caballus, Stephanorhinus cf. hemitoechus, Sus cf. scrofa, Cervus elaphus priscus, Dama dama aff. clactoniana, cf. Hemitragus bonali, large Bos/Bison. Apart from yielding some remains of Homo heidelbergensis and archaeological objects, units TG10–11 of Galería yielded a similar fauna, that however differs in including Megaloceros verticornis dawkinsi? and in having a small Bison? Instead of the large Bos/Bison. Both associations are typical of the middle Middle Pleistocene. A single fossil from the uppermost unit of Sima del Elefante belongs probably to Stephanorhinus hemitoechus.The transition of the Early to Middle Pleistocene coincides with a fundamental change in climate; from this time onward, the 100 ky glacial cycles become a dominant feature. This transition is marked by a series of dispersal events towards Europe, including that of Homo antecessor. The absence of most glacial taxa in the Middle Pleistocene of Spain, as well as the existence of long lineages of “interglacial” taxa within Europe suggests that glaciations had a limited effect on large mammals in Spain and other parts of southern Europe. Homo antecessor may have evolved into Homo heidelbergensis and later into H. neanderthalensis surviving during glaciations in refugia in southern Europe, as did some of the ungulates.
At the lower Miocene locality of Loranca, Spain, numerous skeletal and dental remains of a peculiar Giraffoid have been collected. Many of the long bone diaphyses, especially of the front limbs, possess abnormally thickened multilayered ''pachyostotic'' bone deposits, all adult individuals being affected (MNI = 24). These remains are identified as belonging to a new genus and species of giraffoid Lorancameryx pachyostoticus, close to Teruelia adroveri, another lower Miocene giraffoid from Spain without pachyostosis. In searching for an explanation for the phenomenon of pachyostosis in the Loranca giraffoid, the authors have developed an hypothesis about the evolution of ''abnormal'' bony deposits in artiodactyls, including the appearance of frontal appendages in several Miocene to Recent lineages (Cervidae, Climacoceratidae, Lagomerycidae, Bovidae, Giraffidae, Antilocapridae, Hoplitomerycidae, Palaeomerycidae, Suidae) and of pachyostotic mandibles and maxillae in some Pleistocene Cervidae. The hypothesis is that all these ''abnormal'' bony deposits are simply different strategies for maintaining body/skeletal relations relatively constant in species which undergo marked seasonal body weight fluctuations. The onset of marked seasonality towards the end of the lower Miocene period appears to have been the ''trigger'' for the independent evolution of bony cranial appendages in at least 7 lineages of ruminants and of pachyostotic limb bones in the Loranca giraffoid. Once such appendages had evolved they secondarily took on behavioural significance. The authors also discuss the reasons for the onset of marked seasonality towards the end of the lower Miocene.
This valuable collection of essays presents and evaluates techniques of body-mass estimation and reviews current and potential applications of body-size estimates in paleobiology. Papers discuss explicitly the errors and biases of various regression techniques and predictor variables, and the identification of functionally similar groups of species for improving the accuracy of estimates. At the same time other chapters review and discuss the physiological, ecological, and behavioral correlates of body size in extant mammals; the significance of body-mass distributions in mammalian faunas; and the ecology and evolution of body size in particular paleofaunas. Coverage is particularly detailed for carnivores, primates, and ungulates, but information is also presented on marsupials, rodents, and proboscideans.
To date, fossil deer (Ryukyu jika) Cervus astylodon (Matsumoto, 1926) occurs abundantly from Late Pleistocene fissure and cave deposits in four islands of the Ryukyu Islands : Tokunoshima, Okinawa, Kume, and Ishigaki. This study on Cervus astylodon from the comparative morphological viewpoint reveals that each of the four islands of the Ryukyu Islands yields a characteristic endemic morphotype.
Furthermore, on the island of Kume, several morphotypes can be distinguished. The four Kume morphotypes are characterized by different body-sizes and proportions of teeth and limb bones. These four morphotypes share characteristic metacarpal features, and demonstrate four phases in the dwarfing process. In the process of miniaturization, limb segments distal to the elbow and knee, exhibited a greater degree of dwarfing than did the humerus, femur, and mandibular teeth.
This process also indicates that dwarfism of Cervus astylodon in the Ryukyu Islands occurred bycontinuous morphological change in a relatively short period.
Consequently, morphotypes occur with higher frequency in insular than mainland environments. It is suggested that variation in size and form in deer is more likely to occur in limb bones, especially in metacarpal and metatarsal bones.