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A new species of Eostyloceros (Cervidae, Artiodactyla) from the Late Miocene of the Linxia Basin in Gansu, China
Abstract
A new species, Eostyloceros hezhengensis sp. nov., is established based on a skull with its cranial appendages collected from the Late Miocene Liushu Formation of the Linxia Basin in Gansu Province, northwestern China. It is a large-sized muntjak with a distinct longitudinal ridge along the lateral margin of the frontal bone that joins the antler pedicle. The pedicle is short, cylindrical, robust, and extends posteriorly from the rear of the orbit. The anterior and posterior branches arise from the burr and diverge at an angle of 30 degrees. The posterior branch is relatively long, and its tip is strongly curved posteriorly. The anterior branch is straight and situated anteromedially from the posterior branch. The posterior branch is lateromedially compressed, and the anterior branch has a circular cross section. The morphological observation together with a cladistic and a principal component analysis indicate that E. hezhengensis is more basal than any known species of the genus Eostyloceros in having shorter pedicles, a lower position of the fork above the burr, more slender anterior branches, and a small angle between the anterior and posterior branches. Its age is the middle Late Miocene, corresponding to the late Bahean.
... Zdansky (1925) noticed that this character approaches E. triangularis to modern Muntiacus, however, the poor preservation of the fossil material did not permit to the author to provide a reliable diagnosis for another new genus where E. triangularis probably should be placed. Up to eight species of Eostyloceros were described from the Late Miocene and Pliocene of Eurasia, but not all of them are retained (Thomas 1951;Korotkevich 1970;Vislobokova 1985;Vislobokova 1990;Deng et al. 2014). Thomas (1951) described a new species Eostyloce ros pierensis from the Lower Turolian (MN 11) of the Iberian Peninsula. ...
... Our current knowledge about Asian Eostylo ceros is still incomplete. Thus, the cluster analysis of craniodental characters (including antlers) carried out by Deng et al. (2014) demonstrates that Eostyloceros in its generally accepted understanding is a paraphyletic taxon. ...
... Also worth mentioning here is the relatively short braincase in modern Muntiacus muntjak (the braincase breadth exceeds the braincase length measured as a distance between bregma and inion anatomical points; Croitor, unpublished data) that represents an advanced specialization distinguishing Muntiacus from primitive Miocene cervids with long neurocrania (Vislobokova 1990). The recently discovered cranial material of Eostyloceros hezhengensis is also characterized by a rather short neurocranium (Deng et al. 2014: fig. 3D) as in Muntiacus. ...
We here describe new cervid material from the rich Early Pliocene fauna of Priozernoe
situated on the western slope of the lower part of the valley of Kuchurgan River (Eastern Moldova). An important part of this study aims the taxonomy and systematics of the Kuchurgan cervids. A new genus is established for the small muntjac-like cervids from the Pliocene of Italy, Central and Southeastern Europe. The fauna of Priozernoe contains three cervid species: Praeelaphus australorientalis Croitor, 2017, Procapreolus moldavicus (Janovskaya, 1954), and Eostyloceros pidoplitschkoi Korotkevich, 1964, which we propose to include in a new genus. Such a characteristic association of cervid species resembles the fauna from Priozernoe to Berești (Romania), suggesting the similar geological age. Our results confirm the earlier conclusions on the fauna Priozernoe as the youngest among Kuchurgan faunas and its close affinity with the Moldavian faunal assemblage from the Carbolia Beds.
... In agreement with our hypothesis, a recent phylogenetic analysis 18 also places Eostyloceros blainvillei in an intermediate position between D. elegans and Muntiacus, the only crown cervid considered by the authors, thus preventing any firm conclusion on its crown or stem position. In addition, Deng et al. 44 suggest a paraphyly of the genus Eostyloceros and exclude E. hezhengensis and all the species of Eostyloceros from the crown Cervidae. Despite the results of their phylogenetic analyses, the authors still consider Eostyloceros as a Muntiacinae due to its antler shape 18,44 . ...
... In addition, Deng et al. 44 suggest a paraphyly of the genus Eostyloceros and exclude E. hezhengensis and all the species of Eostyloceros from the crown Cervidae. Despite the results of their phylogenetic analyses, the authors still consider Eostyloceros as a Muntiacinae due to its antler shape 18,44 . Our results confirm the published phylogenetic trees but firmly excludes Eostyloceros from crown Cervidae, and questions the widespread use of antler characteristics in phylogeny. ...
... This character is thus symplesiomorphic for crown deer. More recent phylogenetic analyses mentioned above 18,44 do not seem to confirm the existence of a Muntiacinae clade that includes both Euprox and Eostyloceros, which is here (E. hezhengensis) excluded from Muntiacinae too and even from crown Cervidae. ...
Deer are an iconic group of large mammals that originated in the Early Miocene of Eurasia (ca. 19 Ma). While there is some consensus on key relationships among their members, on the basis of molecular- or morphology-based analyses, or combined approaches, many questions remain, and the bony labyrinth has shown considerable potential for the phylogenetics of this and other groups. Here we examine its shape in 29 species of living and fossil deer using 3D geometric morphometrics and cladistics. We clarify several issues of the origin and evolution of cervids. Our results give new age estimates at different nodes of the tree and provide for the first time a clear distinction of stem and crown Cervidae. We unambiguously attribute the fossil Euprox furcatus (13.8 Ma) to crown Cervidae, pushing back the origin of crown deer to (at least) 4 Ma. Furthermore, we show that Capreolinae are more variable in bony labyrinth shape than Cervinae and confirm for the first time the monophyly of the Old World Capreolinae (including the Chinese water deer Hydropotes) based on morphological characters only. Finally, we provide evidence to support the sister group relationship of Megaloceros giganteus with the fallow deer Dama.
... This situation hampers confident identification and detailed description of the material in question, but contributes to public education and science outreach. Except that the material of Eostyloceros and Promephitis have previously been studied by Deng et al. (2014) and Wang and Qiu (2004), morphological descriptions of the other material are present in this paper. The mandible of H. forstenae was CT scanned, using the 450 kV industrial CT at the Key Laboratory of Vertebrate Evolution and Human Origins, Chinese Academy of Sciences. ...
... The measurements are taken with digital calipers with a precision of 0.1 mm. The morphological nomenclatures and measuring methods follow those of: Qiu, Huang et al. (1987) and Eisenmann et al. (1988) in hipparions, Werdelin (1988) and Werdelin and Solounias (1990) in hyaenids, Guérin (1980) and Deng (2001Deng ( , 2005 in aceratheres, Dong (2008), Wang and Zhang (2011) and Deng et al. (2014) in cervids, and Bohlin (1926) and Ríos et al. (2016) in giraffids. ...
... et sp. indet., and Adcrocuta eximia are reported from Gaojiashan in this paper, and Promephitis hootoni and Eostyloceros hezhengensis were previously reported from this locality (Wang & Qiu 2004;Deng et al. 2014). At least ten species thus occur at the Gaojiashan locality. ...
Two Hipparion species, H. forstenae and H. hippidiodus from Gaojiashan locality in the Linxia Basin, Gansu, China are described in this paper, with the first presence of the former in the Linxia Basin. The lower cheek teeth of Hipparion from Gaojiashan bear deep ectoflexids, and after comparisons to other Chinese Hipparion fossils with deep ectoflexids, the mandibles with deep ectoflexids on the lower premolars suggest their attributions to H. forstenae. The presence of a deep ectoflexid on p2 may be interpretable as an intraspecifically variable feature in H. forstenae, and can also be found in H. dermatorhinum. Besides the Hipparion material, at least 8 species of large mammals occur at the Gaojiashan locality, including Adcrocuta eximia, Chilotherium wimani, Shansirhinus ringstroemi, Eucladoceros cf. proboulei, Palaeotragus cf. coelophrys, Giraffidae gen. et sp. indet., and the previously studied Promephitis hootoni and Eostyloceros hezhengensis. Although the sample from Gaojiashan shares five species with the Yangjiashan fauna, the Gaojiashan fossil assemblage may date to the Baodean because of the presence of H. forstenae. The overall taxonomic composition of the Gaojiashan assemblage at least suggests an approximate Baodean age, which would imply survival of C. wimani into the Baodean of the Linxia Basin.
... In agreement with our hypothesis, a recent phylogenetic analysis 18 also places Eostyloceros blainvillei in an intermediate position between D. elegans and Muntiacus, the only crown cervid considered by the authors, thus preventing any firm conclusion on its crown or stem position. In addition, Deng et al. 44 suggest a paraphyly of the genus Eostyloceros and exclude E. hezhengensis and all the species of Eostyloceros from the crown Cervidae. Despite the results of their phylogenetic analyses, the authors still consider Eostyloceros as a Muntiacinae due to its antler shape 18,44 . ...
... In addition, Deng et al. 44 suggest a paraphyly of the genus Eostyloceros and exclude E. hezhengensis and all the species of Eostyloceros from the crown Cervidae. Despite the results of their phylogenetic analyses, the authors still consider Eostyloceros as a Muntiacinae due to its antler shape 18,44 . Our results confirm the published phylogenetic trees but firmly excludes Eostyloceros from crown Cervidae, and questions the widespread use of antler characteristics in phylogeny. ...
... This character is thus symplesiomorphic for crown deer. More recent phylogenetic analyses mentioned above 18,44 do not seem to confirm the existence of a Muntiacinae clade that includes both Euprox and Eostyloceros, which is here (E. hezhengensis) excluded from Muntiacinae too and even from crown Cervidae. ...
In the last decade, studies based on inner ear morphology flourished thanks to easier access to computer tomography. This structure is deeply associated to the locomotor system and proved to be a source of relevant phylogenetic information. Inner ears in Mesozoic mammals, marsupials, xenathrans, elephants, primates, or rodents are currently under study, while that of ruminants, (one of the most diversified group of living large mammals) remains understudied (Costeur 2014). These analyses usually present classical descriptions, but morphometrics and geometric morphometrics have started to be employed. Unfortunately, the use of geometric morphometrics provides a phenotypic tree and these studies often fail to account for instraspecific variability. Moreover, many discrete morphological characters cannot be taken into account in such analyses. Here we propose to test the phylogenetic relevance of the inner ear morphology based on comparisons of this structure in extinct and extant Cervidae. Stem antlered Cervidae such as Procervulus and Lagomeryx are known as far back as 18 Ma. The relationships within the living deer are stable and the phylogenetic trees resulting from molecular data (mitochondrial and nuclear) are similar. However, no phylogeny based on morphological characters alone reflects the molecular-based results. Moreover, controversial results on divergence time between the groups have been proposed, mainly because of the lack of consensus on the relationships among the fossil cervids. For example, the emblematic Megaloceros is part of the basal radiation of Cervus according to molecular data, of the Dama radiation in combined analyses (molecular and morphological), or even closely related to the “basal Cervini” Eucladoceros when morphological characteristics are taken into account (e.g. Lister et al. 2005). Using a superimposition process (“Landmarks” software), direct comparison between selected inner ears was made. We observed the bony labyrinth of 21 cervid species, including all living tribes and several taxa from key periods: the Early Miocene (origin of stem deer), the Middle Miocene (possible crown deer) and the Plio-Pleistocene (origin of today’s disparity). Intraspecific variability was characterized. Between two specimens (Capreolus capreolus, extant) and six specimens (Procervulus praelucidens, Early Miocene), the differences between juveniles and adults, or intraspecific variability are observed. Similarly to what is observed among the Tragulidae ruminants (Costeur and Mennecart, in prep), no large intraspecific differences are observed. The size of the semicircular canals and the angle between the canals may vary a little, such as the size of the endolymphatic sac. The total cochlea length can also vary by as most as half a turn. However, These differences are smaller than the interspecific variability. Stem deer can easily be distinguished in having the plesiomorphic characters of a large first cochlear turn, a vestibular aqueduct that is aligned with the common crus. The distinction between the living Capreolinae and Cervinae can be made on the basis of the insertion of the lateral semicircurlar canal into the posterior ampula. Cervinae retain the primitive aspect having a high insertion, while in the Capreolinae, the connexion is lateral. This Capreolinae apomorphic state demonstrates that Croizetoceros pyreanicus is the oldest indubitable Capreolinae (6 Ma), even if Late Miocene species have been tentatively attributed to this subfamily (Croitor & Stefaniak 2009). The differences between the various tribes and subtribes can be done based on the shape and position of the endolymphatic sac. Looking at the inner ear morphology, Megaloceros clearly differs from Eucladoceros in having a triangular endolymphatic sac, starting below the end of the common crus, much like in to Dama. On the contrary, Eucladoceros possesses a very elongated endolymphatic sac starting above the common crus. This may be the apomorphy of the Cervus-Rusa lineage that would include “Cervus” ruscinensis as the oldest known representative, dating back to 5 Ma. The general morphological similarities between Megaloceros and Eucladoceros observed in previous phylogenetic analyses are probably linked to symplesiomorphic characteristics and common evolutionary trends in gigantic size and heavy antlers such as proposed by Vislobokova (2013).
... In agreement with our hypothesis, a recent phylogenetic analysis 18 also places Eostyloceros blainvillei in an intermediate position between D. elegans and Muntiacus, the only crown cervid considered by the authors, thus preventing any firm conclusion on its crown or stem position. In addition, Deng et al. 44 suggest a paraphyly of the genus Eostyloceros and exclude E. hezhengensis and all the species of Eostyloceros from the crown Cervidae. Despite the results of their phylogenetic analyses, the authors still consider Eostyloceros as a Muntiacinae due to its antler shape 18,44 . ...
... In addition, Deng et al. 44 suggest a paraphyly of the genus Eostyloceros and exclude E. hezhengensis and all the species of Eostyloceros from the crown Cervidae. Despite the results of their phylogenetic analyses, the authors still consider Eostyloceros as a Muntiacinae due to its antler shape 18,44 . Our results confirm the published phylogenetic trees but firmly excludes Eostyloceros from crown Cervidae, and questions the widespread use of antler characteristics in phylogeny. ...
... This character is thus symplesiomorphic for crown deer. More recent phylogenetic analyses mentioned above 18,44 do not seem to confirm the existence of a Muntiacinae clade that includes both Euprox and Eostyloceros, which is here (E. hezhengensis) excluded from Muntiacinae too and even from crown Cervidae. ...
Modern pecoran ruminants find their origins in the diversified Late Oligocene-Early Miocene hornless taxa. Most of the latter look so primitive and convergent that they never really could be confidently classified, even as stem taxa of the living groups. This lack of agreement has led to problems in understanding the relative phylogenetic positions of the extant families. A still-remaining major issue is the unresolved affinities of the musk-deer family Moschidae. While morphological data tend to favour close ties with cervids, genomic data rather plead for Moschus to be closely related to bovids. The mammalian petrosal bone is known to yield relevant characters for both phylogenetic and functional purposes, but the inner ear embedded within it remains poorly investigated in studies dealing with the evolutionary history of Ruminantia. The inner ear is the organ of hearing and balance. Besides these ecological functions, this structure has been recently shown to bear significant morphological information for phylogeny. The inner ear is virtually unknown in living and fossil ruminants because of its difficult accessibility. Recent advances in non-destructive high-resolution x-ray computer tomography have rendered this organ more accessible. Here we reconstruct the inner ear of all the 21 living ruminant families and tribes together with that of many fossil taxa either attributed to stem groups of the living families or sampled within the Late Oligocene-Early Miocene pecoran radiation. We show that non-pecoran Tragulina have a different inner ear than that of Pecora in having a dorsally branched posterior limb of the lateral semi-circular canal. We trace back the morphology of the moschid inner ear down to the Middle Miocene crown moschidae Micromeryx. This is particularly relevant to the debate on moschid affinities. Fossil stem and crown deers and bovids like Dicrocerus, Heteroprox or Eotragus help understand the evolution of the structure in Pecora. First morphological observations do not clearly support a relationship of Palaeomeryx with Giraffidae. The stem Pecora Prodremotherium, Dremotherium, or Amphitragulus share a junction of the lateral and posterior semi-circular canals. It is distinct from the basal condition of a secondary common crus seen in early artiodactyls. This feature interestingly also occurs in the living deers Cervini and Muntiacini. This study is the first large-scale investigation of the ruminant inner ear including all the living taxa at the tribe to family level and a large set of fossil taxa
... The late Miocene-early Pliocene paleontological record of Eastern Asia has also yielded archaic stem deer that represent a peculiar zoogeographic holdover of the Sino-Malayan zoogeographic zone of that period. The analysis of the inner bony ear of Eostyloceros-a genus containing several small-sized cervid forms with simple two-pointed antlers [17,23,24,46,53,54]-suggest it belongs to an archaic stem deer lineage [55]. ...
... Muntiacus gave a rich diversity of species with body masses ranging from 10-11 kg (M. nanus and M. zhaotongensis [54]) to 40.5 kg (M. gigas [57]). ...
The article describes the paleobiogeographic history of the modern subfamilies so-called "crown deer" of the family Cervidae (Artiodactyla, Mammalia) in the world from the late Miocene to the late Pleistocene. The study overviews the taxonomic diversity and evolutionary radiation of Cervidae from all zoogeographic realms where this systematic group is present in the paleontological record. The evolutionary diversification of the fossil Cervidae is based on the estimations of species body masses that are regarded here as a proxy of occupied ecological niches. The study reveals two important evolutionary radiations of Cervidae during the late Miocene of Eurasia that gave the origin of the modern subfamilies Cervinae and Capreolinae. The evolutionary radiation of Capreolinae during the Pleistocene in South America shows a range of diversity comparable to the late Miocene radiations of Old World deer and provides multiple examples of evolutionary convergences with Eurasian Pleistocene cervids. The article discusses factors that shaped the modern biogeographic distribution of representatives of the subfamilies Cervinae and Capreolinae.
... It is the first time that the species discovered outside Wuxiang. Other species of Eostyloceros found so far in China include E. longchuanensis from Yuanmou in Yunnan Province (Lin et al., 1978) and E. hezhengensis from Hezheng in Gansu Province (Deng et al., 2014); and those abroad include E. pidoplitschkoi from southeastern Moldova and from the Lower Pliocene deposits of the Kuchurgan River at sites Novopetrovka, Yurievka, Voinich in the south of Ukraine, E. propria from the northeastern coast of the Lake Karabastuz in Kazakhstan, E. actauensis from the Middle Miocene of Dzhungarian Aktau in East Kazakhstan, and E. maci from the Pliocene of Olkhon Island in Lake Baikal (Vislobokova, 1990;Deng et al., 2014). The genus is thus distributed mostly in Asia and eastern Europe. ...
... It is the first time that the species discovered outside Wuxiang. Other species of Eostyloceros found so far in China include E. longchuanensis from Yuanmou in Yunnan Province (Lin et al., 1978) and E. hezhengensis from Hezheng in Gansu Province (Deng et al., 2014); and those abroad include E. pidoplitschkoi from southeastern Moldova and from the Lower Pliocene deposits of the Kuchurgan River at sites Novopetrovka, Yurievka, Voinich in the south of Ukraine, E. propria from the northeastern coast of the Lake Karabastuz in Kazakhstan, E. actauensis from the Middle Miocene of Dzhungarian Aktau in East Kazakhstan, and E. maci from the Pliocene of Olkhon Island in Lake Baikal (Vislobokova, 1990;Deng et al., 2014). The genus is thus distributed mostly in Asia and eastern Europe. ...
Five taxa of Cervidae were identified from the new collection at Tuchengzi locality in Huade, Nei Mongol: Eostyloceros blainvillei, E. triangularis, Euprox sp., Cervavitus huadeensis and C. shanxius, among which the first three taxa were discovered for the first time in Huade area, and the dental specimens of the last two taxa from Tuchengzi locality are also described for the first time. E. blainvillei is a large muntiacine, diagnosed by long, thick and medially curved main beam and a relatively long brow tine emerging directly from the burr. It was originally found in Yushe Basin in Shanxi Province, and reported later from Qaidam Basin in Qinghai Province. Huade is the third area yielding the species. E. triangularis distinguishes from E. blainvillei by its special main beam with triangular cross sections, and it was reported only from Yushe Basin. Its presence in Huade extended its geographic distribution northward to Nei Mongol. Euprox is a transitional form of cervids from permanent antlers to seasonally deciduous ones and it is found in several localities across Eurasia. Huade is the third area in Nei Mongol yielding the taxon after Tung Gur and Siziwang Qi (Siziwang Banner). C. huadeensis is a pliocervine with four tines, and particularly with two distal tines sword-like. It is found at Tuchengzi for the second time, but not elsewhere so far, and it seems to be an endemic taxon. C. shanxius distinguishes from C. novorossiae by absence of Palaeomeryx fold on lower molars. It is widely found in Shanxi, Shaanxi, Gansu and Nei Mongol of northern China. It is often found with numerous specimens in a locality that indicates it lived in large herds. E. blainvillei, E. triangularis and C. shanxius are main members of Yushe I, i.e. Mahui Formation or Baodean. Euprox appeared also mainly in the Late Miocene. The geological age of Tuchengzi locality based on excavated cervids is therefore the Late Miocene. The numerous specimens of C. shanxius indicates forested environment in Huade area in that age.
... Located at the northeastern border of the Tibetan Plateau, the Linxia Basin ( Fig. 1) is one of the most fossiliferous late Cenozoic basins known in China, with rich discoveries from the Late Oligocene to the Early Pleistocene (Deng, 2004b;Deng et al., 2004b;Deng, 2009;Deng et al., 2013b). Hundreds of fossil species have been recognised in this basin, including many new species Deng and Qiu, 2007;Qiu et al., 2007;Wang and Deng, 2011;Deng et al., 2013b;Deng et al., 2013c;Deng et al., 2014a;Deng et al., 2014b;Hou and Deng, 2014;Qiu et al., 2014;Wang et al., 2015;Shi and Deng, 2021). The Linxia Basin also provides records of one of the most continuous mammalian evolutionary history, with four large faunas, i.e., Late Oligocene Paraceratherium fauna; Middle Miocene Platybelodon fauna, which can be further subdivided into three faunas, from Early/Middle Miocene, middle Middle Miocene, and late Middle Miocene, respectively; Late Miocene to Pliocene hipparion fauna, which can be further subdivided into six faunas, including four successive faunas from early Late Miocene to latest Late Miocene, one from Miocene/Pliocene boundary, and one from Late Pliocene; and Early Pleistocene Equus fauna (Deng et al., 2013b and this study). ...
The Linxia Basin has produced some of the most complete and fossiliferous fossil assemblages among the known
late Cenozoic basins including diverse carnivoran elements. In this work, we summarize the chronological
framework for its fossil Carnivora based on current knowledge. A total of 80 species of Carnivora, belonging to 55
genera and 10 families, are currently recorded in 10 assemblages. Three faunas with Carnivora of Middle
Miocene age, include amphicyonids, hemicyonids, and percrocutids. Five assemblages dating to the late Miocene
and the earliest Pliocene hipparion faunas, are represented by very abundant ictitherine hyaenids, the giant
hyaena Dinocrocuta, the large machairodont Amphimachairodus and diverse metailurine felids, and various
mustelids. The Late Pliocene records scant material, but the Early Pleistocene Equus fauna is well represented by
canids, hyaenids, a lynx, and Megantereon. A major Carnivoran guild turnover occurs at the boundary of the
Middle and Late Miocene. The Linxia Basin experienced earlier aridification in the Late Miocene than did Europe,
likely correlated with the rise of Tibet. The medial body size of Carnivora remains largely unchanged from the
Middle Miocene to the Early Pleistocene. Crown-group members (at the family level) increased dramatically in
proportion during the Late Miocene, and remained largely unchanged since the latest Miocene, suggesting that
the Late Miocene is the major period of emplacement of current Carnivora phylogenetic structure.
... The decline of late Miocene giraffids in Southern Asia coincided with the evolutionary radiation of plesiometacarpal cervids (subfamily Cervinae) in Southeast Asia. The Late Miocene cervine radiation evolved along with stem cervid lineages such as Eostyloceros (Deng et al., 2014;Mennecart et al., 2017) and holometacarpal deer (Dong, 2011). ...
This article presents a description of new antler remains from five fossiliferous sites (Sardhok, Panjan Sher Shahana, Puran, Jari Kas, and Potha) of the Upper Siwaliks in Pakistan. The systematic study of the antler material revealed the presence of six cervid forms: Metacervocerus punjabiensis, Rucervus sp., Panolia sp., Hyelaphus sp., Praesinomegaceros bakri, and a poorly represented large cervid that shows a certain affinity with “Eucladoceros sp.” from the Early Pleistocene of Kuruksai (Tajikistan). The remains of Panolia represent the earliest known paleontological record of this cervid lineage. Unlike Metacervocerus and Rucervus that have had phylogenetically closely related counterparts in east and north of the Alpine-Himalayan mountain belt, the evolution of Panolia took place in the Indian subcontinent. The entry of Panolia lineage into the Indian subcontinent marks its phylogenetic split from the main Cervus/Rusa evolutionary branch. The earliest dispersal events of cervids into the Indian subcontinent was preceded by the late Miocene evolutionary radiation and ecological diversification of the subfamily Cervinae in Southeastern Asia. Praesinomegaceros and Metacervocerus most probably entered the Indian subcontinent via Central Asia. Possibly, this is also the case of “Eucladoceros sp.” from Kuruksai (Tajikistan). The dispersal ways of Panolia and Rucervus remain unclear. The dispersals of small-sized cervids (Muntiacus and Hyelaphus) into the Indian subcontinent was triggered by the establishment of the 100-ky glaciation cycle during the Middle Pleistocene: the sea level dropped during glacial peaks and opened to them the dispersal route from Sundaland in the South.
... Euprox (Hou, 2015), Eostyloceros (Deng, Wang, Shi, Li, & Li, 2014) and their direct ancestors, for example, Dremotherium ) and therefore seems to be a plesiomorphic character. The large preorbital depression in the small Pudu could be a plesiomorphic condition, not affected by the secondary size reduction. ...
In Ruminantia, the lacrimal bone forms a considerable part of the facial skeleton, and the morphology of its facial facet is highly variable when compared to other mammals. In this study, we quantify the species‐specific variability in size and shape of the lacrimal facial facet in species of Cervidae (deer) and relate it to systematics and various aspects of their ecology and behavior. We sampled 143 skull specimens from 10 genera; 12 Moschus and 3 Tragulus specimens were used as outgroups. We find that size and shape of the lacrimal facial facet allow differentiating most species analyzed here, except for Mazama gouazoubira and Capreolus capreolus. Size and shape of the lacrimal facial facet vary widely across Cervidae regardless of their systematic relationships, ecology or behavior. Thus, we could not detect a unique signature of adaptational criteria in lacrimal morphology. Our data indicate that the lacrimal facial facet scales allometrically with skull size, in particular, the lacrimojugal length scales positively and the lacrimomaxillar length scales negatively. However, correlation analyses did not reveal any differences in the integration of the lacrimal bone with any specific skull module in any of the species compared. Lastly, we could not ascertain any correlation between the size and position of the preorbital depression with the size and shape of the lacrimal facial facet. We conclude that the lacrimal facial facet is highly flexible and may rapidly adjust to its surrounding bones. Its allometric growth appears to be an example of exaptation: changes in size and shape in the context of the increase of the skull length provide lacrimal contacts, in particular, a lacrimojugal one, which may serve to reduce mechanical loads resulting from increasingly larger antlers in large cervids.
... Euprox (Hou, 2015), Eostyloceros (Deng, Wang, Shi, Li, & Li, 2014) and their direct ancestors, for example, Dremotherium ) and therefore seems to be a plesiomorphic character. The large preorbital depression in the small Pudu could be a plesiomorphic condition, not affected by the secondary size reduction. ...
Most mammals possess a lacrimal bone, which builds the rostral rim of the orbita. In deer (Cervidae, Artiodactyla), the bone acquires considerable size compared with other mammals. It is characterized by a depression hosting the lacrimal gland, whose secretion is used for territory marking. The lacrimal bone and its depression are very variable in both size and shape within cervids being useful for systematics. However, the underlying causes for this variability remain so far unexplained.
Given the central position of the lacrimal bone in the facial skull (it articulates with the nasal, frontal, sphenoid, jugal, and maxilla), it is likely that the bone reflects adaptations to different ecological niches and different behaviour. This hypothesis coincides with the evolution of cervids, where early small forest dwellers from tropical /subtropical habitats grew in size and adapted to temperate and subarctic regions.
In our presentation, we give an overview of size and shape in some living cervid species and the correlation with behaviour and habitat. We use the results to infer on behaviour and ecology in fossil cervids.
The site of Çandir has yielded one of the best collections of Giraffidae from the Middle Miocene of the Eastern Mediterranean. It is here referred to a single new species, whose comparison with other contemporaneous remains from this area and the Siwaliks raises some interesting phylogenetic problems.
There are six Ruminant species at Çandir, not including the Giraffid Giraffokeryx. The smallest one is a Micromeryx, a genus left as Eupecora incertae sedis, but its determination at species level must await full revision of the genus. A large rare form is provisionally referred to Palaeomeryx. Also rare is a new species of the primitive cervid Heteroprox, with antlers similar to the European ones, but with quite a different dentition. Bovids are much more common, with the hypsodont genera Turcocerus and Hypsodontus, and the more generalised Tethytragus. The ruminants provide no evidence for putting Çandir later than Paşalar, but they reveal ecological differences, Çandir Locality 3 being especially open.
The discovery by Birger Bohlin of a series of vertebrate fossil sites in the twin lakes region (Tuosu Nor and Keluke Nor) of eastern Qaidam Basin during the Sino-Swedish Expedition in 1931 and 1932 was a major milestone in vertebrate paleontology for the Tibetan Plateau. Qaidam fossil mammals collected by Bohlin still represent the best collections from the plateau and serve as an important reference point for a period of time that saw dramatic climatic changes. The more than 4600 m of strata in eastern Qaidam, spanning over 13 million years in time, are ideal for establishing a detailed biostratigraphic record, but Bohlin’s published specimens lack any reference to stratigraphy, which causes much confusion about the nature of his “Tsaidam Fauna.”
Bohlin did, however, make a fairly detailed documentation of locality information, much of which remains buried in archival records in Stockholm. This paper is an attempt to reconstruct Bohlin’s fossil localities by synthesizing relevant archives and historical accounts, as well as field observations of our own during the past 13 years. Fieldnotes in Swedish are translated to English and several field sketches are published for the first time. As a result, we are able to relocate many fossil localities that are of stratigraphic and taxonomic importance.
Bohlin mainly collected in three major areas, along the northwestern shore of Tuosu Nor, south of Huaitoutala, and south of Quanshuiliang train station, each with stratigraphic settings of their own. In his published descriptions, all fossils are included in his “Tsaidam Fauna.” With the help of his notes and sketches and our field verification, we are now in a position to recognize at least three faunal horizons among Bohlin’s collections: middle Miocene (Tunggurian) Olongbuluk Fauna, early late Miocene (early Bahean) Tuosu Fauna, and early Pliocene (Yushean) Huaitoutala Fauna.
Of these fossil sites, Bohlin’s Tuosu Nor locality is the best constrained within the modern stratigraphic framework due to the relatively short section present along the northern bank of Tuosu Nor. His fossil localities are confined to a narrow band of about 100 meters in total thickness. Bohlin’s Huaitoutala (Ulan-utsur) localities are, however, more scattered both spatially and stratigraphically. Fortunately, only a few localities produced fossils of biochronologic significance, and most are low in the section, probably belonging to the Olongbuluk Fauna.
The majority of Bohlin’s Qaidam collections comes from what he informally called the “general strips,” which is not far from the present day Quanshuiliang railway station. Approximately 250 m of strata extend laterally for several kilometers. Almost all endemic bovids that Bohlin described, such as Tsaidamotherium, Olonbulukia, Qurliqnoria, Tossunnoria, are produced from this area. The Quanshuiliang region also produces some of the earliest occurrences of Hipparion in China. Preliminary correlation of the Quanshuiliang section with an existing magnetic section in Huaitoutala suggests that the local first appearance of Hipparion is close to the boundary between chrons C5r.1r and C5r.1n, about 11.1 Ma.
Rescue excavations carried out from 2007 to 2010 at the Shuitangba lignite field in Zhaotong, Yunnan Province, not only resulted in the discovery of a new hominoid cranium and the addition of a new hominoid site in Yunnan, but the finding of a new muntjak with following characters: the brow tine is very close to the burr, which is moderately developed; the main beam extends backward and somewhat laterally, and then turns somewhat medially; and the neocrista and entocingulum are developed on the upper molars. The four limbs are relatively short. A cladistic analysis shows that the new materials represent a muntjak that is not in a sister-group relationship with Muntiacus leilaoensis from Yuanmou Late Miocene hominoid site, but rather represents an independent branch. Muntiacus zhaotongensis sp. nov. is proposed for the new muntjak materials. "Metacervulus sp." from the "Yongle lignite field" recovered in 1978 is a synonym of the new species. The so-called "Yongle lignite field" is actually the Shuitangba lignite field. The layers yielding the materials of M. zhaotongensis sp. nov. are all within C3An.1n based on paleomagnetic correlation, with an estimated age between 6.1 to 5.9 Ma (terminal Miocene).
This chapter describes the Late Cenozoic biostratigraphy of the Linxia Basin in northwestern China. The Linxia Basin harbors a variety of Late Cenozoic fossils of different plants and animals as well as land mammals. Reptiles are represented mainly by a great number of specimens of Testudo from the late Miocene red clay deposits, which is an important member of the Hipparion fauna. After sketching the geological setting, magnetostratigraphy, and biochronology of the Linxia Basin, the chapter analyzes the abundant mammalian fossils from the Linxia Basin, with particular emphasis on their exact localities and horizons. By doing so, it is able to update the faunal lists from specific localities, establish lithological and biostratigraphic criteria for correlation, and conduct paleomagnetic studies of key sections of the area.
A new genus and species of deer, Lucentia iberica, is described from the lower Turolian (MN 11) locality of Crevillente 2 (Alicante province, Spain). The two-tined Lucentia is not a representative of Muntiacinae. From the emplacement on the cranial cavity and moderate backward inclination of the pedicles, and from the monopodial construction of the antlers, Lucentia is probably the sister group of the clade Cervinae + Odocoileinae. The hypothesis that Lucentia is most closely related to Cervinae is poorly supported by dental features. The Cervinae-Odocoileinae split must have occurred during the late Miocene and it being later, during the Pliocene, when the different lineages acquired the lateral metacarpal reduction, either plesiometacarpal or telemetacarpal states. -from Authors
An Eostyloceros antler fragment is described. The specimen makes part of the Montopoli (Lower Valdarno, Tuscany) collection preserved in the Geo-Paleontological section of the Natural History Museum of Florence. This site is the type locality of the homonymous Faunal Unit of the Italian Biochronological scheme; it includes the latest Early Villafranchian Mammal Age faunas. The occurrence of an Eostyloceros species among this material however, is in disagreement with the age of the Montopoli local fauna, which has been correlated with the Gauss-Matuyama boundary and thus dated to about 2.6 Ma. The youngest remains of this genus come from sites of eastern Europe Early Pliocene in age (Ruscinian Mammal Age). The possibility that the antler is an external element to the Montopoli assemblage is therefore contemplated.
The Yuanmou Basin of Yunnan Province is a hotspot for the research of early hominid evolution and its environmental background.
During the implementation of the Program “Research of the Origin and Evolution of Early Man and Its Environmental Background”
from 1998 to 2001, a huge quantity of hominoid and mammal fossils were collected. The present report systematically describes
anEuprox robustus sp. nov. identified during the recent laboratory work on the collected material.Euprox is a group of earliest cervids with true antlers. The new species is the third one of the genus discovered in China. Judged
by its morphologic characteristics, the new species feed on juicy and tender leaves of dicotyledon. It implies that the vegetation
of its epoch in the Yuanmou Basin is a kind of southern subtropical evergreen forest and the climate is humid and temperate
with evident seasonality. The latter is mostly influenced by the monsoon and secondly by the latitude.
Keywords
Euprox
-Cervidae-Artiodactyla-Yuanmou-Late Miocene
In this paper a new chilothere species, Acerorhinus hezhengensis represented by associated skull and lower jaw from Late Miocene deposits of Hezheng, Gansu, is described. The Chinese 'Diceratherium' problem is reviewed and analysed. Systematically, the Chinese 'Diceratherium' has scarcely any direct relation with the true Diceratherium of North America. Heissig's referral of the Chinese 'Diceratherium' to Chilotherium as a subgenus is here disputed. A series of distinctive characters warrant the Chinese 'Diceratherium' a generic rank, for which Acerorhinus created by Kretzoi in 1942 is adopted. Acerorhinus shares common apomorphic features exclusively with Chilotherium. They may represent a separate tribe in the subfamily Acerotheriinae: Chilotherini.- Authors
The extended excavations from 1998 to 2000 in Yuanmou, Yunnan Province, southwestern China, resulted in the discovery of a new Late Miocene (ca 9–7 Ma) hominoid locality with associated mammalian fauna at Leilao. It includes a new muntjak, Muntiacus leilaoensis sp. nov. It has similar morphology to the extant forms and is the earliest record of the Muntiacus. This discovery shows that the Muntiacus are a persistent and conservative group in contrast with the other rapidly evolving cervids since ca 9–7 Ma.
Recent expeditions in the northeastern Qinghai-Tibetan Plateau have produced additional mammalian fossils from the Lower Baodean (equivalent to Vallesian, Late Miocene), and, significantly, a new large and the earliest muntjac, Muntiacus noringenensis sp. nov. It is phylogenetically closer to the Late Pliocene M. fenghoensis and extant M. vuqangensis than the other fossil and extant muntjacs are. The existence of this muntjac and other folivores in the Late Miocene Qaidam Basin suggests a forested period in the basin and the evolution from a forested and humid environment to a desert one today. Such a dramatic evolution in environments is an evidence of the effect of the uplift of the Qinghai-Tibetan Plateau and of its influence on continental climates.
Qaidam Basin is the largest terrestrial basin of the Tibetan Plateau and has the most continuous sedimentary record in the Cenozoic. Although constituting the first known fossil vertebrates of the Tibetan Plateau and discovered in early scientific expeditions in the 1930s, mammalian fossils from Qaidam Basin have not played a significant role in studying the basin stratigraphy. Our explorations in the Qaidam Basin during the past 8 years have resulted in the first well documented collection of fossil vertebrates from there. In addition to being the best organisms for terrestrial biostratigraphy in the Cenozoic, the new fossil vertebrates offer a unique window into the ancient biotas that chronicle the late Cenozoic uplift of the plateau and associated environmental changes.
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Apd burr
Lmd burr
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