The phylogeny of the Rhinocerotoidea (Mammalia, Perissodactyla)

A new phylogeny of the Superfamily Rhinocerotoidea is proposed, based upon an analysis of shared derived characters of the skull, teeth and skeleton. Hyrachyus is considered the primitive sister-taxon of the three rhinocerotoid families (Amynodontidae, Hyracodontidae, Rhinocerotidae), and the amynodonts appear to be the sister-group of hyracodonts and rhinocerotids. The relationships of primitive hyracodonts and rhinocerotids are clarified. Menoceras and Pleuroceras (new Subfamily Menoceratinae) are removed from the Diceratheriinae, since they appear to be more closely related to higher rhinoceroses than they are to Diceratherium. Of the three major monophyletic groups of higher rhinocerotids (aceratherines, teleoceratines and rhinocerotines), the last two groups are more closely related to each other than either is to aceratherines. Toxotherium and Schizotheroides are tentatively removed from the Rhinocerotoidea and placed in the Lophiodontidae.     

Use of mitochondrial DNA sequences to test the Ceratomorpha (Perissodactyla:Mammalia) hypothesis

Wood (J. Mamm. 18, 106–118; 1937) united the superfamilies Tapiroidea sensu stricto and Rhinocerotoidea in the suborder Ceratomorpha and aligned the Ceratomorpha with the suborder Hippomorpha within the order Perissodactyla. Although the monophyly of the Ceratomorpha appears now well-supported in paleontological and morphological analyses, the molecular relationship among the three extant superfamilies Tapiroidea, Rhinocerotoidea, and Equoidea has not yet been examined due to the limited amount of molecular information on tapirs. In the present study, we examined the phylogenetic position of Tapiroidea, represented by the complete mitochondrial cytochrome b gene (1140 bp) of a lowland tapir ( Tapirus terrestris ), and a Indian tapir ( Tapirus indicus ), relative to modern horses, zebras, donkeys, and rhinoceroses. The phylogenetic analyses using standard parsimony, neighbour-joining and maximum likelihood algorithms revealed monophyly of the Perissodactyla and three clearly distinct lineages: the modern horses, tapirs, and rhinoceroses. However, the sister-taxon relationship of the tapirs to either the rhinoceroses or the horses was not resolved conclusively in the bootstrap analysis. Spectral analysis, in which phylogenetic information is displayed independently of any selected tree, revealed that the DNA sequences available do not contain enough phylogenetic signal for any of the alternative hypotheses on the basal diversification of perissodactyls. The short branch lengths among the three perissodactyl lineages suggest that they diverged within a relatively short period, a finding consistent with molecular divergence datings and the fossil evidence that indicates a major radiation of the early perissodactyls approximately 54–50 million years ago.     

Anatomy and ultrastructure of the proboscis in Mesorhynchus terminostylis (Platyhelminthes,Rhabdocoela)

The ultrastructural organization of the proboscis in Mesorhynchus terminostylis is distinctly different from that in other members of the Polycystididae in which it is currently classified. The sheath epithelium is formed by three belts, all with intra-epithelial nuclei. The apical belt of the bipartite cone epithelium has a single intrabulbar nucleus, and the basal belt possesses five insunk nucleiferous cell parts behind the bulb. Six types of glands surface through the epithelia; the three types emerging through the cone epithelium can be homologized with those described for Polycistis naegelii. Only uniciliary receptors are found in the epithelium. The musculature in the bulb has a very loose appearance, and the bulbar septum appears to be a bipartite basement membrane. The septum can be considered the basement membrane of the cone epithelium as if the contractile portion of the inner longitudinal muscles have invaded the epithelium and come to lie between the epithelial cells and the basement membrane. Thus the inner musculature of the bulb is entirely intraepithelial as is the case for Psammorhynchus tubulipenis and Cytocystis clitellatus. The systematic position of M. terminostylisremains uncertain but seems to lie between Psammorhynchus and Cytocystis on one hand and Koinocystididae and Polycystididae on the other.     

Evolutionary patterns of Perissodactyla and Artiodactyla (Mammalia) with different types of digestion

Described and discussed are the comparative and functional anatomy of the digestive tract in ungulates, as well as the retention of digesta in the total digestive tract and in its compartments. Additionally, the history of distribution of ungulates under special consideration of the connections between continents and the development of different biomes is discussed. The last 40 million years are considered. During the Caenozoic the forestomach-fermenting Artiodactyla were strongly radiating, which is in contrast to conditions in hindgut-fermenting Artiodactyla (Suidae) and Perissodactyla (Equidae, Rhinocerotidae, Tapiridae). The Pecora have been especially successful forestomach-fer-menters because they are able to reduce particle size of digesta through the process of rumination and thus increase digesta passage. This is advantageous because not only the metabolic products (mainly volatile fatty acids) of microbial activity are made available to the mammal, but also a higher percentage of those nutrients that can be autoenzymatically digested by the ungulate.     

The entotympanic of Equus caballus (Perissodactyla,Mammalia)

In the majority of extant placental mammals the bulla tympanica is composed of two skeletal elements, the entotympanic and the ectotympanic. Former studies revealed that the presence of an entotympanic in the bulla tympanica of extant Perissodactyla is restricted to Rhinocerotidae. The existence of the entotympanic in Tapiridae and Equidae remained speculative. Here we present the first evidence of an entotympanic, strictly speaking rostral entotympanic, in the domestic horse, Equus caballus. The enchondrally ossified entotympanic can be easily separated from the desmal ectotympanic by its greater thickness and by its cancellous bone texture in a late fetal stage. Both elements are separated by a suture that is in the process of coalescence. The complete fusion of the two elements and the unification of bone texture are almost accomplished at birth but the entotympanic and ectotympanic assume the same thickness obviously not until early postnatal development. Based on modern phylogenetic hypotheses we can conclude that the common ancestor of Perissodactyla must have possessed a well-developed entotympanic, probably only evident in their fetal life. This must be considered as a plesiomorphic character state of this order, because the entotympanic is a neomorphic apomorphy of placental mammals. However, the prenatal fusion of the entotympanic and the ectotympanic is an apomorphy of Equus caballus and possibly of the Equidae as a whole.     

Asteropeia and Physena (Caryophyllales): A case study in comparative wood anatomy

Previous analyses ofAsteropeia andPhysena have not compared the wood anatomy of these genera to those of Caryophyllales s.l. Molecular evidence shows that the two genera from a clade that is a sister group of the core Caryophyllales. Synapomorphies of theAsteropeia-Physena clade include small circular alternate pits on vessels, presence of vasicentric tracheids plus fiber-tracheids, presence of abaxial-confluent plus diffuse axial parenchyma, and presence of predominantly uniseriate rays. These features are analyzed with respect to habit and ecology of the two genera. Solitary vessels, present in both genera, are related to the presence of vasicentric tracheids. Autapomorphies in the two genera seem related to adaptations byPhysena as a shrub of moderately dry habitats (e.g., narrower vessel elements, abundant vasicentric tracheids, square to erect cells in rays) as compared to alternate character expressions that seem related to the arboreal habit and humid forest ecology ofAsteropeia. The functional significance of vasicentric tracheids and fiber-tracheids in dicotyledons is briefly reviewed in the light of wood anatomy of the two genera.     

Phylogeny of the Ferungulata (Mammalia: Laurasiatheria) as determined from phylogenomic data

Great progress has been made toward resolving the evolutionary relationships among extant mammals, yet there are still areas of disagreement. The relationships among ferungulates that have high quality draft genome sequences available (i.e. dog, cow, horse) are unresolved, and thus we examined their phylogeny using currently known mammalian 1:1 orthologs. This dataset consists of 40 million base pairs from 2705 protein-coding genes. Maximum likelihood and Bayesian analyses of the combined and individual gene phylogenies strongly support a sister grouping of cow and horse to the exclusion of dog although topology tests could not rule out a horse and dog sister group relationship.     

A new elasmothere (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China

Ningxiatherium euryrhinus sp. nov. is a relatively large elasmothere rhino from the Linxia Basin in northwestern China found in the early Bahean, which corresponds to the early Vallesian (MN 9), ca. 11.1 Ma. It is much larger than the extant Ceratotherium simum and has a single horn. Ningxiatherium is similar to the late Miocene Parelasmotherium from Gansu and Shanxi, but differs in having partially ossified nasal septum, terminal nasal horn boss, shallower nasal notch above the P3/P4 boundary, and much shorter premolars. N. euryrhinus sp. nov. differs from N. longirhinus from the late Miocene of Zhongning, Ningxia in northwestern China by its larger size, relatively wider nasals, shallower nasal notch, and sub-quadrangular occlusal surface of M3. In the Linxia Basin, the other early late Miocene (Bahean or Vallesian) elasmotheres are known from complete skulls or isolated teeth. They include two genera and three species: Parelasmotherium linxiaense, P. simplum, and Iranotherium morgani. They lack a nasal septum, but Ningxiatherium has one. The age of N. euryrhinus is early late Miocene based on direct association with biochronologic indicator taxa, such as Dinocrocuta gigantea, Hipparion dongxiangense, and Chilotherium primigenius. N. euryrhinus is more primitive than N. longirhinus sp. nov. in having a sub-quadrangular M3, a shallower nasal notch, and the presence of DP1 in adults.     

Identification of a new contingency-based response in honey bees (Apis mellifera) through revision of the proboscis extension conditioning paradigm

The two experiments reported present new information in the area of classical conditioning experiments with honey bees. Experiment 1 establishes a single unconditioned stimulus (US) technique as a preferred technique for conditioning of the proboscis extension response. Experiment 1 further identifies a new head turn response which occurs when the standard compound US technique is used. Experiment 2 demonstrates that the newly identified head turn response is contingency-based and provides important new response to the repertoire of honey bee learning experiments.     

Skull evolution in the rhinocerotidae (Mammalia,Perissodactyla): Cartesian transformations and functional interpretations

Cartesian transformation, applied as a landmark morphometric method, is used to investigate some of the evolutionary shape changes leading to the skulls of the modern rhinoceroses. The early Oligocene genusSubhyracodon serves as the primitive shape from which the extant genera (Dicerorhinus, Rhinocerso, Diceros, andCeratotherium) have been transformed. Coordinate data for 21 landmarks, defined in lateral view, are analyzed by the computer program Thinplate Splines. Each of the four transformations are interpreted separately as shape change fromSubhyracodon. Computed results forRhinoceros are also compared with previous results obtained by visual interpretation (the classic method). Among the extant genera,Ceratotherium andRhinoceros have the most derived shapes and are opposites with respect to orientation of the occiput and relative size of the mandible angle. The significance of these foci of change is discussed in terms of the functions of the masseter and posterior temporalis muscles. In head positions associated with feeding on short vs. tall grasses, the two skull shapes are consistent with a role for these muscles in support of a large mandible against gravity. This common factor may help to explain both shapes.     

Sussemionus, a new subgenus of Equus (Perissodactyla,Mammalia)

The new subgenus of Equus, Sussemionus, is defined by peculiar dental characters so far unknown, or exceptional in late Pleistocene and extant Equus; it was in consequence assumed to be restricted to the early and middle Pleistocene. During that period, it was highly successful, ranging from North America to Ethiopia, and included dry-adapted (E. granatensis-like) and more humid-adapted (E. coliemensis-like) species. Recent molecular and osteological analyses concurred to prove its survival until ca 45 KYBP in Khakassia, southwest Siberia, Russia.     

Flies and concealed nectar sources: morphological innovations in the proboscis of Bombyliidae (Diptera)

Bee-flies (Bombyliidae) have morphological adaptations of the mouthparts to particular floral traits. To investigate this the short, plesiomorphic proboscis of Hemipenthes morio was compared with the long, apomorphic proboscis of Bombylius major . A novel feeding position enables B. major to use flowers that open to the side as additional nectar sources. The new horizontal feeding position is enabled by the prolonged ventral base of the proboscis. Bombylius major exploits deep corolla tubes with an elongate proboscis, and an increased efficiency in both the suction pumps and the sealing mechanisms of the proboscis. The exploitation of narrow corolla tubes is made possible by the shift from a sponging feeding mode, exhibited by H. morio , to the exclusively sucking mode in B. major . Besides quantitative changes in the proportions of the different proboscis components, labellar movements as well as the structures of saliva distribution are changed along with this shift. The labial musculature of B. major does not significantly differ from the plesiomorphic state, since both examined species do not only feed on nectar, but also on pollen.     

山西天镇辛窑子早更新世犀科化石新材料（英文）

20世纪80年代在桑干河盆地一带考察泥河湾层时,在山西省天镇县南高崖乡的辛窑子沟一带发现了很多哺乳动物化石地点并出土了大量哺乳动物化石。最近对其中犀科化石的研究表明在辛窑子沟一带的早更新世地层中产出的犀类化石至少有两个种,裴氏板齿犀(Elasmotheriumpeii)和泥河湾披毛犀(Coelodontanihowanensis),可能还存在过第三个种。由于标本保存状况不理想,暂时鉴定为梅氏犀相似种(Stephanorhinus cf. S. kirchbergensis)。虽然后者的形态大小与梅氏犀最接近,但和泥河湾披毛犀也有相似之处,因此也有可能是这两个种之一的种内变异类型。同样,产于下沙沟地点被德日进和皮维托鉴定为有疑问的中国犀(Rhinoceros sinensis (?))也可能是梅氏犀或泥河湾披毛犀的种内变异类型。迄今为止在广义泥河湾盆地发现的早更新世犀类有两个确定的种和两个不确定的种。出土裴氏板齿犀的地点为下沙沟、山神庙咀、大黑沟及辛窑子沟;出土泥河湾披毛犀的地点为下沙沟、大南沟、东谷坨、山神庙咀及辛窑子沟。有疑问的中国犀仅出现在下沙沟,梅氏犀相似种仅出现在辛窑子沟。     

Rhinocerotidae (Mammalia,Perissodactyla) from the middle Pleistocene levels of Grotta Romanelli (Lecce,southern Italy)

The rhinoceros remains collected during the past century in the lower levels XII (= K) and XI (= I) of the famous Pleistocene locality of Grotta Romanelli (Lecce, southern Italy) are described and compared in detail for the first time. Some remains are referred to Stephanorhinus sp. and others are assigned here to the late early-middle Pleistocene European species Stephanorhinus hundsheimensis based on several morphological characters. Based on its olivine-bearing texture, the volcanoclastic ash sampled from some rhinoceros bones can be referred to the first phase of the Monte Vulture activity (around 630 ka). The results of the stable isotope analyses suggest that the climate in the lowest levels of Grotta Romanelli could have been more arid than it was at the time of the upper level IX, which is generally referred to the late Pleistocene. In addition, both recent day δ¹⁸O_ppt values and MAT are very similar to values calculated for levels X and XII, suggesting that the climate at those times may have been close to the Present one, whereas climate in level IX may have been somewhat cooler. The presence of Stephanorhinus hundsheimensis suggests a middle Pleistocene age for the lower levels of Grotta Romanelli, in agreement with the results obtained from the volcanoclastic material.     

The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its palaeobiological implications

The Pliocene Beds of the Pisco Formation on the southern coast of Peru have yielded three new skulls of the walrus-like odontocete, Odobenocetops (Delphinoidea, Cetacea). Two of the skulls are from a slightly younger horizon than the holotype of O. peruvianus and belong to a different species, O. leptodon . The holotype of O. leptodon bears a 135 cm long needle-like right tusk and a small 25 cm long left tusk, the apex of which was erupted. The third skull, referred to a female of O. peruvianus , bears two small tusks similar in size to the small left tusk of the holotype. The new specimens include periotic, tympanic and ear ossicles, which confirm referral to the Order Cetacea and the morphology of the humerus reinforces affinities of odobenocetopsids to monodontids. Because the anterodorsal edge of its orbit is slightly concave O. leptodon had reduced anterodorsal binocular vision, a condition compensated for by the probable presence of a small melon (and inferred echolocation). The head was bent ventrally when swimming in such a way that the long tusk was approximately parallel to the axis of the body. The extremely salient occipital condyles of Odobenocetops are indicative of great mobility of the neck, probably related to bottom-feeding. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 423–452.     

Cranial anatomy of the Paleocene plesiadapiform Carpolestes simpsoni (Mammalia, Primates) using ultra high-resolution X-ray computed tomography, and the relationships of plesiadapiforms to Euprimates

Central to issues surrounding the origin of euprimates, affinities of Paleocene Carpolestidae have been controversial. Carpolestids have been classified as plesiadapoid primates, tarsiiform euprimates, dermopterans, or the sister taxon of euprimates to the exclusion of other plesiadapiforms, based exclusively on dental or postcranial data. Newly discovered crania of Carpolestes simpsoni from the latest Paleocene of the Clarks Fork Basin, Wyoming, are the first described for the family Carpolestidae. The two best preserved skulls were studied using ultra high-resolution X-ray computed tomography. Comparison of these specimens to those of other stem primates (Plesiadapiformes) demonstrates that the diversity of cranial morphology in this group is greater than previously thought. Carpolestes differs from euprimates and is similar to other plesiadapiforms (Ignacius and Plesiadapis) in lacking a postorbital bar and having a relatively long rostrum. Carpolestes is similar to fossil euprimates and Plesiadapis in having a bullar morphology consistent with a petrosal origin, and differs from Ignacius, in which the bulla is composed of the entotympanic. Carpolestes differs from primitive euprimates and all other known plesiadapiforms in possessing a two-chambered auditory bulla, similar to that of modern Tarsius. However, Carpolestes had an internal carotid artery (ICA) that took a transpromontorial route from a posteromedially positioned posterior carotid foramen (pcf), unlike Tarsius, in which this artery takes a perbullar route from an anterolaterally positioned pcf. Carpolestes has clear grooves on the promontorium for both the promontorial and stapedial arteries, indicating that it had an unreduced internal carotid circulation, similar to that of early euprimates. Carpolestes differs from primitive euprimates and some specimens of Ignacius in not having bony tubes surrounding the branches of the ICA. Cladistic analysis of cranial data fails to support a close relationship of Carpolestidae to either tarsiiform euprimates or extant Dermoptera, but suggests a close relationship between Carpolestidae, Plesiadapidae, and Euprimates.