A primitive ceratomorph (Perissodactyla, Mammalia) from the early Tertiary of Europe

Hyracotherium cuniculus Owen, 1842, from Palaeocene–Eocene boundary strata in NW Europe, was formerly considered to be an equoid. It is here placed in Cymbalophus gen. nov. , in the perissodactyl suborder Ceralomorpha, which comprises tapiroids and rhinocerotoids, as one of its most primitive and earliest members. This revised classification is based on an evaluation of its advanced versus primitive character states, and comparison with a variety of other primitive perissodactyls. Cheek tooth occlusal relationships in the early stages of ceratomorph evolution are discussed and contrasted with the fundamental perissodactyl morphotype as represented in Hyracotherium. It is suggested that the Lophiodontidae (formerly considered to be ceratomorphs; may be more closely related to chalicotheres; that the Isectolophidae may be the sister-group of chalicotheres and lophiodonts plus ceratomorplis; and that Cymbalophus is the most primitive known member of the newly restricted Ceratomorpha. Cymbalophus is placed tentatively in the family Helaletidae.     

Chromosome evolution in perissodactyla

Comparative painting has provided a wealth of useful information and helped to reconstruct the pathways of karyotype evolution within major eutherian phylogenetic clades. New data have come from gene localizations, BAC mapping and high throughout sequencing projects that enrich and provide new details of genome evolution. Extensive research on perissodactyl genomes has revealed not only increased rates of chromosomal rearrangements, but also an exceptionally high number of centromere repositioning events in equids. Here were combined new physical mapping, comparative painting and genome sequencing data to refine the putative ancestral karyotype maps and to revise the previously proposed scenario of perissodactyl karyotype evolution.     

Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls

Anthracobunidae is an Eocene family of large mammals from south Asia that is commonly considered to be part of the radiation that gave rise to elephants (proboscideans) and sea cows (sirenians). We describe a new collection of anthracobunid fossils from Middle Eocene rocks of Indo-Pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships, instead implying that they are stem perissodactyls. Cranial, dental, and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus. Analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land, but spent a considerable amount of time near water. This new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins.     

Phylogenetics of Perissodactyla and Tests of the Molecular Clock

Two mitochondrial genes, the protein-coding cytochrome c oxidase subunit II (COII) gene and a portion of the 12S rRNA gene, were used for phylogenetic investigation of the mammalian order Perissodactyla. The primary objective of the study was to utilize the extensive fossil record of perissodactyls for calibrating molecular clocks and comparing estimates of divergence times using both genes and two fossil calibration points. Secondary objectives included clarification of previously unresolved relationships within Tapiridae and comparison of the results of separate and combined analyses of two genes. Analyses included several perissodactyl lineages representing all three families (Tapiridae, Equidae, and Rhinocerotidae), most extant genera, all four species of tapirs, two to four species of rhinoceros, and two species of Equus. The application of a relatively recent fossil calibration point and a relatively ancient calibration point produced greatly different estimates of evolutionary rates and divergence times for both genes, even though a relative rates test did not find significant rate differences among taxa. A likelihood-ratio test, however, rejected a molecular clock for both genes. Neither calibration point produced estimates of divergence times consistent with paleontological evidence over a range of perissodactyl radiations. The combined analysis of both genes produces a well-resolved phylogeny with Perissodactyla that conforms to traditional views of interfamilial relationships and supports monophyly of neotropical tapirs. Combining the data sets increases support for most nodes but decreases the support for a neotropical tapir clade because the COII and 12S rRNA data sets are in conflict for tapir relationships. Received: 6 January 1999 / Accepted: 2 August 1999     

日本九州西部赤崎组和中甑组始新世哺乳动物:初步结果及其与亚洲陆相哺乳动物分期的对比(英文)

日本已知时代最早的古近纪哺乳动物化石产自九州西部熊本县天草(Amakusa)地区始新世地层赤崎(Akasaki)组以及鹿儿岛县甑岛(Koshiki Islands)相当层位的中甑(Nakakoshiki)组。初步研究显示这两个组产出的哺乳动物分异度较高,包括9个目的至少18种动物。赤崎动物群有3种trogosine裂齿类、2种冠齿兽科全齿类、1种等脊貘科奇蹄类、2种dichobunoid偶蹄类、1种下齿兽科踝节类、1种西瓦兔猴科灵长类、1种未定食虫类以及2种可能的梳趾鼠类。中甑动物群包括1种冠齿兽科全齿类、2种小型雷兽科奇蹄类、1种高冠的下齿兽科踝节类、2种啮齿类和1种细齿兽科食肉类。这两个日本哺乳动物组合很接近早-中始新世界线。由于出现了trogosine裂齿类和雷兽,它们的时代要晚于伯姆巴动物群,极有可能与亚洲大陆阿山头期哺乳动物群相当。这两个动物群还包含了在亚洲大陆没有发现过的几个新种,并具有独特的哺乳动物组合。     

Perissodactyl diversities and responses to climate changes as reflected by dental homogeneity during the Cenozoic in Asia

Cenozoic mammal evolution and faunal turnover are considered to have been influenced and triggered by global climate change. Teeth of large terrestrial ungulates are reliable proxies to trace long‐term climatic changes due to their morphological and physicochemical properties; however, the role of premolar molarization in ungulate evolution and related climatic change has rarely been investigated. Recently, three patterns of premolar molarization among perissodactyls have been recognized: endoprotocrista‐derived hypocone (type I); paraconule–protocone separation (type II); and metaconule‐derived pseudohypocone (type III). These three patterns of premolar molarization play an important role in perissodactyl diversity coupled with global climate change during the Cenozoic in Asia. Those groups with a relatively higher degree of premolar molarization, initiated by the formation of the hypocone, survived into Neogene, whereas those with a lesser degree of molarization, initiated by the deformation of existing ridges and cusps, went extinct by the end of the Oligocene. In addition, the hypothesis of the “Ulan Gochu Decline” is proposed here to designate the most conspicuous decrease of perissodactyl diversity that occurred in the latest middle Eocene rather than at the Eocene–Oligocene transition in Asia, as conventionally thought; this event was likely comparable to the contemporaneous post‐Uintan decline of the North American land fauna.     

Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla)

The systematics and taxonomy of early Eocene equids are investigated. A paraphyletic sequence of equid taxa is recovered from a phylogenetic analysis of 40 taxa and 121 characters. This analysis supports the identification of Hyracotherium as a primitive equoid and its restriction to the genotype, Hyracotherium leporinum . Sifrhippus gen. nov. is erected for the sister taxon of all other equids, Hyracotherium sandrae Gingerich. Minippus gen. nov., the next more-derived equid clade, is erected for two small equids, M. index Cope and M. jicarillai nov. species. Arenahippus gen. nov. is erected for the next three sequentially more-derived equid taxa, A. grangeri Kitts, A. aemulor Gingerich, and A. pernix Marsh. The genus Xenicohippus , which is the next more-derived equid clade, is redefined to include X. craspedotum Cope. Eohippus Marsh, the next more-derived equid taxon, is resurrected for E. angustidens Cope. Pliolophus , the only early Eocene equid from Europe, is identified as the sister taxon to Protorohippus , a sequence of successively more-derived equid taxa consisting of P. montanum Wortman and P. venticolum Cope. Protorohippus venticolum is identified as the sister taxon of Orohippus . Systemodon Cope is resurrected for S. tapirinum Cope. This taxon has historically been placed within Hyracotherium yet this analysis allies it with Cymbalophus near the base of the perissodactyl radiation. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 134 , 141–256.     

New localities of Late Eocene vertebrate footprints from the Tarom Mountains,Northwestern Iran

We document Late Eocene vertebrate footprints from the Tarom Mountains of Iran that represent a significant addition to the record of proboscidean and perissodactyl footprints. These footprints are from sandstones and tuffaceous sandstones of strata equivalent to the Kond Formation that overlie middle Eocene sedimentary rocks and are overlain by Oligocene volcanics. The footprints are preserved at 16 tracksites from 10 distinct stratigraphic levels. The mammal footprints include the oldest known proboscidean tracks, assigned to Proboscipeda enigmatica Panin & Avram and to cf. Proboscipeda isp. Evident perissodactyl tracks are common, tridactyl footprints with distinct digit shapes and proportions assigned to the new ichnogenus and ichnospecies Moropopus elongatus. Footprints of small, hopping, rodent-like mammals are identified as Musaltipes taromi new ichnospecies. Other mammal footprints from the Tarom tracksites are indeterminate, and bird footprints are assigned to Avipeda isp. The Tarom tracksites document the oldest record of proboscidean footprints, and this indicates that proboscideans had reached the northern shore of Tethys by Late Eocene time. The abundance and distinctiveness of the Tarom perissodactyl tracks mirrors the abundance and diversity of moropomorph perissodactyls during the Middle-Late Eocene. The Tarom tracksites are the most extensive record of Eocene vertebrate footprints known from Iran.

http://zoobank.org/urn:lsid:zoobank.org:pub:F430F9DB-728F-4162-9791-DC2A5CC5ED43     

Bird and Mammal Footprints From the Tertiary of Navarre (Western Pyrenees)

A rich variety of vertebrate footprints is known from a number of Upper Eocene to Lower Miocene localities of Navarre (western Pyrenees). The sediments were deposited in a wide range of depositional environments, from marginal marine to diversified terrestrial. Abundant bird tracks have been found in the coastal deposits of the Upper Eocene Liedena Sandstone of the Yesa and Itzagaondoa areas. Ciconiiformes-like (Leptoptilostipus pyrenaicus) and Charadriiformes-like (Charadriipeda ichnospp.) footprints have been recognized. Mammal ichnites have been discovered in the Oligocene and Lower Miocene deposits of Navarre. Equoid perissodactyl ichnites similar to those of Plagiolophustipus occur in the Oligocene fluviatile rocks of the Mués Sandstone of Olexoa and the Rocaforte Sandstone near Oibar and Sada. Trackways of entelodontids (Entelodontipus) are known in fluviatile-palustrine beds of the Oligocene Mués Sandstone of Olkotz. Additionally, bird (Charadriiformes-like) tracks are known in fluviatile-palustrine floodplain deposits of the Lower Miocene Ujué Formation of Los Arcos. In the same area, the Desoio and Los Arcos outcrops have also yielded perissodactyl trackways of possible Equoidea. Trackways of rhinocerotids (?) and artiodactyls (possibly Pecoripeda) are described from the Lower Miocene (Ramblian) palustrine limestones marginal to the Lerín Formation of Kaparroso and from alluvial fan deposits of the Uncastillo-Perdón Formation of Altzorritz, respectively.     

The Evolution of Mammalian Olfactory Receptor Genes

We performed a comparative study of four subfamilies of olfactory receptor genes first identified in the dog to assess changes in the gene family during mammalian evolution, and to begin linking the dog genetic map to that of humans. The human subfamilies were localized to chromosomes 7, 11, and 19. The two subfamilies that were tightly linked in the dog genome were also tightly linked in the human genome. The four subfamilies were compared in human (primate), horse (perissodactyl), and a variety of artiodactyls and carnivores. Some changes in gene number were detected, but overall subfamily size appeared to have been established before the divergence of these mammals 60-100 million years ago.     

Mammalian Male Mutation Bias: Impacts of Generation Time and Regional Variation in Substitution Rates

In mammals, males undergo a greater number of germline cell divisions compared with females. Thus, the male germline accumulates more DNA replication errors, which result in male mutation bias—a higher mutation rate for males than for females. The phenomenon of male mutation bias has been investigated mostly for rodents and primates, however, it has not been studied in detail for other mammalian orders. Here we sequenced and analyzed five introns of three genes (DBX/DBY, UTX/UTY, and ZFX/ZFY) homologous between X and Y chromosomes in several species of perissodactyls (horses and rhinos) and of primates. Male mutation bias was evident: substitution rate was higher for a Y chromosome intron than for its X chromosome homologue for all five intron pairs studied. Substitution rates varied regionally among introns sequenced on the same chromosome and this variation influenced male mutation bias inferred from each intron pair. Interestingly, we observed a positive correlation in substitution rates between homologous X and homologous Y introns as well as between orthologous primate and perissodactyl introns. The male-to-female mutation rate ratio estimated from concatenated sequences of five perissodactyl introns was 3.88 (95% CI = 2.90–6.07). Using the data generated here and estimates available in the literature, we compared male mutation bias among several mammalian orders. We conclude that male mutation bias is significantly higher for organisms with long generation times (primates, perissodactyls, and felids) than for organisms with short generation times (e.g., rodents) since the former undergo a greater number of male germline cell divisions. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Deborah Charlesworth]     

Observations on rhinoceros tongue morphology

The rhinoceros tongue is distinguished among perissodactyl organs by its sagittally divided intermolar eminence crowned by a dextral and a sinistral cluster of circumvallate papillae. These structures were originally noted in the Indian rhinoceros by Owen (1852) but have since remained unmentioned for this, or for any other rhinoceros form. They are here described for specimens of the Indian, African White and African Black rhinoceroses. The structural and functional nature of the intermolar eminence is established and this entity is distinguished from an hitherto unrecognized topographical component of the tongue, designated the pars elliptica linguae. The lingual morphology of the three rhinoceros forms is summarised and certain intergeneric differences are noted.     

A NEW PRIMITIVE CHALICOTHERE (PERISSODACTYLA,MAMMALIA) FROM THE EARLY EOCENE OF HUBEI, CHINA

'I.DESCR1PTIoNorderPerissodactylaOwen,l848SuperfamilyCbalicotherioideacall,l872'Family?EomoropidaeVret,l958GehusDanjiangiagen.nov.TypespeciesDaniiangiapingigen.etsp-nov.DiagnoaisSma1lprimitivecha1icothere(lengthofM/l=8.3mm),skullshallowwithcranialrostrumrelatively1ong,mandibularhorizontalramusandmandibularsymphysisfairlylongandsha1low,teeth1ow-crowned,dentitionformula:3.l.4.3/3.1.4.3.Premolarnonmolariform,P2/2tIenchant,P4/withincipientmesostyle.M/widerthan1engthwithW-shapedectoIoph,…     

Anthracotheriid artiodactyl Anthracokeryx and an upper Eocene age for the Youganwo Formation of southern China

Abstract

The two recently collected mammalian dentary fragments from the Eocene Youganwo Formation of Guangdong Province, southern China, are referred to the anthracotheriid species Anthracokeryx naduongensis based on phylogenetic analysis and size comparison. One of these specimens (SYSU-M-1) is the first mammal fossil described from the Youganwo Formation. It was attributed previously to the perissodactyl genus Lunania. Anthracokeryx naduongensis was described originally from the lower upper Eocene Na Duong Formation in Northern Vietnam. The second record of this species supports a basal upper Eocene correlation for the Youganwo Formation, which was estimated previously as middle or late Eocene.     

新疆准噶尔盆地三个泉地区几种始新世哺乳类

本文记述了准噶尔盆地北缘三个泉地区依希白拉组中采到的踝节类、蹠行类、全齿类和恐角类化石。根据这些化石,依希白拉组的时代可能和北美中始新世勃力吉期相当。文内,通过文献分析,认为周明镇和胡长康(1956)记述的真恐角兽的前臼齿可能产自三个泉附近的依希白拉组。     

The complete mitochondrial DNA sequence of the greater Indian rhinoceros, Rhinoceros unicornis, and the Phylogenetic relationship among Carnivora, Perissodactyla, and Artiodactyla (+ Cetacea)

The sequence (16,829 nt) of the complete mitochondrial genome of the greater Indian rhinoceros, Rhinoceros unicornis, was determined. Like other perissodactyls studied (horse and donkey) the rhinoceros demonstrates length variation (heteroplasmy) associated with different numbers of repetitive motifs in the control region. The 16,829-nt variety of the molecule includes 36 identical control region motifs. The evolution of individual peptide-coding genes was examined by comparison with a distantly related perissodactyl, the horse, and the relationships among the orders Carnivora, Perissodactyla, and Artiodactyla (+ Cetacea) were examined on the basis of concatenated sequences of 12 mitochondrial peptide-coding genes. The phylogenetic analyses grouped Carnivora, Perissodactyla, and Artiodactyla (+ Cetacea) into a superordinal clade and within this clade a sister group relationship was recognized between Carnivora and Perissodactyla to the exclusion of Artiodactyla (+ Cetacea) . On the basis of the molecular difference between the rhinoceros and the horse and by applying as a reference to Artiodactyl/Cetacean divergence set at 60 million years ago (MYA), the evolutionary divergence between the families Rhinocerotidae and Equidae was dated to approximately 50 MYA.      

云南中甸新发现的早更新世哺乳动物

描述了最近发现于横断山脉中段的一批哺乳类化石 ,具体地点位于云南迪庆州中甸县城 (香格里拉 )以南约 6km处的公路旁。这批材料包括灵长类 (Procynocephaluscf.P .wimani)、啮齿类 (Hystrixsp .)、食肉类、奇蹄类 (Equusyunnanensis)及偶蹄类 ,其中以鹿类 (Cer vavitusultimus、Eucladocerossp .、Cervus (Rusa)yunnanensis等 )和牛羊类最为丰富 ,揭示了当时一种混合的森林 -草原环境。从动物群总体面貌来看 ,这批哺乳类可与元谋人动物群相比 ,时代为更新世早期。但其中有些种类与北方早更新世类群相似 ,如真枝角鹿及羚羊等 ,表明该动物群也与北方动物群之间存在着某种生物地理上的联系。