Molecular phylogeny of the tribe Bovini (Mammalia: Artiodactyla): alternative placement of the Anoa

Noncoding regions from the genes encoding aromatase cytochrome P450 and lactoferrin have been sequenced in ten bovine and one cervid species for an investigation of the evolutionary relationships within the tribe Bovini. The evolutionary rate of DNA-nucleotide alterations along the ancestral bovine lineage amounts to 0.38% per million years, as estimated from this combined 0.478-kb-single copy nuclear (scn) DNA sequence data set. Whereas rate homogenity is apparent within the Bovini, the relative rate test suggests that the boselaphine lineage (as represented by Boselaphus) has evolved at only about one third of the rate found within the Bovini. Consistent with other results, the scnDNA data provide evidence for (i) a monophyletic origin of the Bovini, (ii) a sister group position of the Boselaphini, and (iii) two different clades within the Bovini, the buffaloes (Bubalus and Syncerus) and the cattle (Bos/Bibos and Bison). Surprisingly, the results indicate very clearly that the enigmatic dwarf buffalo of Sulawesi Island (Anoa depressicornis) is most closely related to Boselaphus and that the divergence from the true Bovini occurred close to the base of bovine cladogenesis in the Middle Miocene (≈ 14—12 million years ago).     

Phylogenetic analysis of the tribe Bovini using microsatellites

The objective of the present study was to determine if the generally accepted phylogenetic relationships in the tribe Bovini correspond to a phylogenetic scheme derived from polymorphisms at 20 bovine microsatellite loci. This study comprises 17 representative populations: eight Bos taurus, two Bos indicus, one Poëphagus, one Bibos, one Bison, three Bubalus and one Syncerus. Phylogenetic analyses using (δμ)² and chord (D_C) distances revealed substantial divergence among species. Neighbor‐joining trees with both distance measures showed only minor differences. Bos taurus and Bos indicus grouped first, followed by Bos frontalis (Mithan) and Bos grunniens (Yak), Bison bison branched off next and Bubalus bubalis and Syncerus caffer emerged as the two most divergent species from the Bos clade. These findings would suggest that Bos, Poëphagus, and Bibos should be integrated into the Bos genus with each group classified as a subgenus. On the other hand, Bison, Bubalus and Syncerus should each be considered a separate genus. Direct estimates of the divergence times were calculated using the (δμ)² genetic distance. Bos taurus and Bos indicus were estimated to have diverged 0·31–0·82 MYA, Bos and Poëphagus: 0·57–1·53 MYA, Bos and Bibos: 0·57–1·52 MYA, Bos and Bison: 0·46–1·23 MYA, Bos and Bubalus: 1·85–4·93 MYA and Bos and Syncerus: 0·98–2·61 MYA.     

Les Lagomerycidae (Artiodactyla, Mammalia) de FranceThe French Lagomerycidae (Cervoidea, Artiodactyla, Mammalia)

The Lagomerycidae, a small family of the Cervoidea, is known in France by the genera Lagomeryx, Ligeromeryx and Stephanocemas. Lagomeryx contains three species, the other two genera one species only. This family is known from MN3 to MN5.     

Postcranial morphology and locomotion of the Eocene raoellid Indohyus (Artiodactyla: Mammalia)

Raoellids are small, raccoon-sized Eocene artiodactyls, closely related to archaic cetaceans (archaeocetes) that have poor representation of postcranial elements in the fossil record. Little is known of the aquatic and terrestrial locomotor affinities of raoellids due to the paucity of their fossil record, leaving a critical gap in our understanding of the earliest portion of the artiodactyl marine invasion. To address this gap, a comparative morphological analysis of the postcranial elements was undertaken based on newly recovered elements of the raoellid Indohyus, archaeocetes and extant artiodactyls. Greater than 200 postcranial elements of Indohyus were described, and some limb element cross-sections were visualised via paleohistological thin sections and CT scans. Results show that during terrestrial locomotion, Indohyus probably had a digitigrade posture and mediolaterally stabilised limbs that functioned mostly in flexion and extension within the parasagittal plane. Quantification of midshaft cross-sectional area for some elements of Indohyus showed an osteosclerotic cortex, a skeletal characteristic associated with aquatic behaviours. Indohyus may represent a critical intermediate in the evolution of the cetacean terrestrial-to-aquatic body plan, as it bears gracile postcranial element proportions similar to a terrestrial artiodactyl but also an incipient form of osteosclerosis compared to pakicetid archaeocetes.     

Phylogeny and classification of tribe Aedini (Diptera: Culicidae)

The phylogeny and classification of tribe Aedini are delineated based on a cladistic analysis of 336 characters from eggs, fourth‐instar larvae, pupae, adult females and males, and immature stage habitat coded for 270 exemplar species, including an outgroup of four species from different non‐aedine genera. Analyses of the data set with all multistate characters treated as unordered under implied weights, implemented by TNT version 1.1, with values of the concavity constant K ranging from 7 to 12 each produced a single most parsimonious cladogram (MPC). The MPCs obtained with K values of 7–9 were identical, and that for K = 10 differed only in small changes in the relationships within one subclade. Because values of K < 7 and > 10 produced large changes in the relationships among the taxa, the stability of relationships exemplified by the MPC obtained from the K = 9 analysis is used to interpret the phylogeny and classification of Aedini. Clade support was assessed using parsimony jackknife and symmetric resampling. Overall, the results reinforce the patterns of relationships obtained previously despite differences in the taxa and characters included in the analyses. With two exceptions, all of the groups represented by two or more species were once again recovered as monophyletic taxa. Thus, the monophyly of the following genera and subgenera is corroborated: Aedes, Albuginosus, Armigeres (and its two subgenera), Ayurakitia, Bothaella, Bruceharrisonius, Christophersiomyia, Collessius (and its two subgenera), Dahliana, Danielsia, Dobrotworskyius, Downsiomyia, Edwardsaedes, Finlaya, Georgecraigius (and its two subgenera), Eretmapodites, Geoskusea, Gilesius, Haemagogus (and its two subgenera), Heizmannia (and subgenus Heizmannia), Hopkinsius (and its two subgenera), Howardina, Hulecoeteomyia, Jarnellius, Kenknightia, Lorrainea, Macleaya, Mucidus (and its two subgenera), Neomelaniconion, Ochlerotatus (subgenera Chrysoconops, Culicelsa, Gilesia, Pholeomyia, Protoculex, Rusticoidus and Pseudoskusea), Opifex, Paraedes, Patmarksia, Phagomyia, Pseudarmigeres, Rhinoskusea, Psorophora (and its three subgenera), Rampamyia, Scutomyia, Stegomyia, Tanakaius, Udaya, Vansomerenis, Verrallina (and subgenera Harbachius and Neomacleaya), Zavortinkius and Zeugnomyia. In addition, the monophyly of Tewarius, newly added to the data set, is confirmed. Heizmannia (Mattinglyia) and Verrallina (Verrallina) were found to be paraphyletic with respect to Heizmannia (Heizmannia) and Verrallina (Neomacleaya), respectively. The analyses were repeated with the 14 characters derived from length measurements treated as ordered. Although somewhat different patterns of relationships among the genera and subgenera were found, all were recovered as monophyletic taxa with the sole exception of Dendroskusea stat. nov. Fifteen additional genera, three of which are new, and 12 additional subgenera, 11 of which are new, are proposed for monophyletic clades, and a few lineages represented by a single species, based on tree topology, the principle of equivalent rank, branch support and the number and nature of the characters that support the branches. Acartomyia stat. nov. , Aedimorphus stat. nov. , Cancraedes stat. nov. , Cornetius stat. nov. , Geoskusea stat. nov. , Levua stat. nov. , Lewnielsenius stat. nov. , Rhinoskusea stat. nov. and Sallumia stat. nov., which were previously recognized as subgenera of various genera, are elevated to generic status. Catageiomyia stat. nov. and Polyleptiomyia stat. nov. are resurrected from synonymy with Aedimorphus, and Catatassomyia stat. nov. and Dendroskusea stat. nov. are resurrected from synonymy with Diceromyia. Bifidistylus gen. nov. (type species: Aedes lamborni Edwards) and Elpeytonius gen. nov. (type species: Ochlerotatus apicoannulatus Edwards) are described as new for species previously included in Aedes (Aedimorphus), and Petermattinglyius gen. nov. (type species: Aedes iyengari Edwards) and Pe. (Aglaonotus) subgen. nov. (type species: Aedes whartoni Mattingly) are described as new for species previously included in Aedes (Diceromyia). Four additional subgenera are recognized for species of Ochlerotatus, including Oc. (Culicada) stat. nov. (type species: Culex canadensis Theobald), Oc. (Juppius) subgen. nov. (type species: Grabhamia caballa Theobald), Oc. (Lepidokeneon) subgen. nov. (type species: Aedes spilotus Marks) and Oc. (Woodius) subgen. nov. (type species: Aedes intrudens Dyar), and seven are proposed for species of Stegomyia: St. (Actinothrix) subgen. nov. (type species: Stegomyia edwardsi Barraud), St. (Bohartius) subgen. nov. (type species: Aedes pandani Stone), St. (Heteraspidion) subgen. nov. (type species: Stegomyia annandalei Theobald), St. (Huangmyia) subgen. nov. (type species: Stegomyia mediopunctata Theobald), St. (Mukwaya) subgen. nov. (type species: Stegomyia simpsoni Theobald), St. (Xyele) subgen. nov. (type species: Stegomyia desmotes Giles) and St. (Zoromorphus) subgen. nov. (type species: Aedes futunae Belkin). Due to the unavailability of specimens for study, many species of Stegomyia are without subgeneric placement. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species.     

Phylogenetic relationships in the tribe Oxyptilini (Lepidoptera,Pterophoridae, Pterophorinae) based on morphological data of adults

The monophyly of the tribe Oxyptilini and phylogenetic relationships of the genera embraced in this tribe were examined using 171 (75 binary and 96 multistate) characters of adult morphology. The study material included 98 species of 30 genera, representing all previously recognized genera of Oxyptilini, together with the genera Sphenarches, Antarches, Diacrotricha, and Cosmoclostis, four species of Oidaematophorini, three species of Platyptiliini, as well as three and two other species belonging to Pterophorini and Exelastini respectively. Two Agdistis species were used as outgroups. The cladistic analysis resulted in six equally parsimonious trees. A majority of the recovered synapomorphic characters have previously been used in the taxonomy of the subfamily. However, 25 novel characters were found. The monophyly of Oxyptilini was supported, although only with homoplastic characters and low amounts of tree confidence; the genera Capperia, Procapperia, Paracapperia, Oxyptilus, Megalorhipida, and Trichoptilus were found to be nonmonophyletic; Sphenarches and Antarches were recovered as members of Oxyptilini; the two genera Cosmoclostis and Diacrotricha were placed out of Oxyptilini, inside the tribe Pterophorini; and close affinity of the genus Dejongia to Stangeia, Stenodacma, Megalorhipida, Trichoptilus, and Buckleria species was revealed. Four new combinations, Cosmoclostis lanceata (Arenberger) comb. nov. , Nippoptilia regulus (Meyrick) comb. nov. , Capperia tadzhica (Zagulajev) comb. nov. , and Buckleria negotiosus (Meyrick) comb. nov. are proposed; Capperia insomnis Townsend was considered as a senior synonym of Procapperia hackeri Arenberger syn. nov. , Buckleria negotiosus (Meyrick) as a senior synonym of Buckleria vanderwolfi Gielis syn. nov. , and Oxyptilus variegatus Meyrick syn. nov. as a junior synonym of Oxyptilus secutor Meyrick. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 484–547.     

A New Species of Amphirhagatherium (Choeropotamidae, Artiodactyla, Mammalia) from the Late Eocene Headon Hill Formation of Southern England and Phylogeny of Endemic European 'anthracotherioids'

A new species of artiodactyl, Amphirhagatherium edwardsi sp. nov., is described from the Late Eocene (Priabonian) Headon Hill Formation of the Hampshire Basin, southern England. The Haplobunodontidae, in which Amphirhagatherium is usually placed, has recently been combined with the monotypic Choeropotamidae, both essentially European endemic families. New anatomical information is forthcoming from both the new species and recently published data on related species. A cladistic analysis of taxa included in the two families, the possible anthracotheriid Thaumastognathus and the enigmatic Tapirulus , was conducted to test the relationships implied by observed morphological similarities. The genus Anthracobunodon is shown to be paraphyletic and is here synonymized with Amphirhagatherium . Choeropotamus and Thaumastognathus are sister taxa nested with three species of Haplobunodon and Haplobunodon is paraphyletic and polyphyletic, but this clade is too weakly resolved internally for reliable taxonomic changes. Lophiobunodon Tapirulus are sister taxa nested with a fourth species of Haplobunodon . The synonymy of the Haplobunodontidae with the Choeropotamidae is upheld and close relationship of the family with the Anthracotheriidae is argued to be unlikely. Choeropotamids are inferred to have had mixed frugivorous and browsing herbivorous diets. They seem to have diversified in the northern parts of Europe, some terminal taxa having originated following southward dispersal.     

牛科(哺乳纲:偶蹄目)动物与食物有关的适应形态模式

利用逐步分辨分析方法（Stepwise discriminant analysis,SCDA）检测了广义牛科动物的颅齿部结构,这些结构特征可以作为采食行为生态适应特征。在本研究中,测量了72种广义牛科动物的28个颅齿部结构。逐步分辨分析方法得出了6种采食方式适应类型：一般粗食者、新鲜禾草粗食者、开阔生境混合型采食者、精食者、郁闭生境混合型采食者、食果者。用103个标本检测了分辨指标的预测能力,所用标本为缺损标本,大多数缺少一项或多项结构。从这些标本获得的分辨函数的平均预测能力为94％,比用72种广义牛科动物标本建立的分辨函数的平均预测能力（98％）低一些。从一个颅齿部结构小样本建立的分辨函数可以用于考古发掘物中不完整标本的研究。这些指标与用颅下结构测量建立的运动能力和生境选择的指标相结合,可以推断古牛科动物的个体生态学以及古环境重建。     

Phylogenetic analysis and classification of fossil and recent Alcelaphini Mammalia: Bovidae

Cladistic principles are applied to a group of antelope species, three-quarters of which are extinct. Only cranial and mandibular characters are used. The present phylogenetic hypothesis suggests diat the Alcelaphini are monophyletic; that the genera Parmularius and Damaliscus share a recent common ancestry; and that die extant Hunter's hartebeest should not be removed from Damaliscus into a separate genus Beatragus as several authors have done. A re-evaluation is suggested of die systematic position of Lichtenstein's hartebeest, as being phylogenetically closer to me wildebeests, genus Connochaetes , than to the type species of the hartebeest genus Alcelaphus. Some problems arising in the application of cladistic principles to such a low ranking group, treating die fossil majority exactly equivalently with recent taxa, are discussed. Comments on recency of common ancestry as it affects hybridization potential in bovids, on speciation and extinction rates arise.     

Tachyoryctes makooka (Tachyoryctini,Spalacidae, Rodentia) and its bearing on the phylogeny of the Tachyoryctini

Abstract: The Rhizomyinae is a subfamily of myomorph rodents within the family Spalacidae. It is subdivided into two tribes: the Asian Rhizomyini and the Tachyoryctini. Even though the origin of the Tachyoryctini is to be found in Asia, they are nowadays restricted to Africa. African Tachyoryctini are known from the Late Miocene and include a single genus with five species: the recently discovered Tachyoryctes makooka, which is described in detail in this work, T. pliocaenicus, T. konjiti and the two extant T. splendens and T. macrocephalus. Their closest Asian counterparts are the Late Miocene Protachyoryctes and Eicooryctes. A cladistic analysis involving all the above‐mentioned taxa was carried out. Protachyoryctes tatroti turned out to be the most basal species of the ingroup followed by the Ethiopian T. makooka. Both taxa show some primitive traits like the protosinus on the first upper molars, short but distinct mesolophids on the lower molars and the presence of the mure on the cheek teeth. T. pliocaenicus and the most derived Tachyoryctini share the synapomorphic reduction or loss of these characters. The increase in hypsodonty evidenced in this lineage is correlated with the transition from a humid to a drier climate that started at the beginning of the Pliocene.     

Phylogeny and evolutionary history of the Aplodontoidea (Mammalia: Rodentia)

Athough over a hundred species of fossil aplodontoids have been described since the extant species, Aplodontia rufa (the mountain beaver), was first described by Rafinesque in 1817, a thorough survey of the relationships among all the species in this clade has not been undertaken since McGrew's study in 1941. Here, a complete phylogenetic analysis of all published species of aplodontoids is used to reconstruct the evolutionary relationships within the clade, and to present an updated classification of the Aplodontoidea. Several of the traditionally recognized subfamilies are found to be paraphyletic, namely the Prosciurinae, the Allomyinae, and the Meniscomyinae. Others, however, including the Aplodontinae and the Mylagaulidae, appear to be monophyletic. These latter two taxa, which include all of the hypsodont members of the aplodontoid clade, seem to be sister taxa. The history of the aplodontoid clade shows several episodes of rapid diversification in the Early Oligocene, the Late Oligocene, and the Early to Middle Miocene. The Ansomyinae and Aplodontinae show comparatively low speciation rates. The patterns of change in morphology and evolutionary rates suggest a need for a more detailed study of the causes of diversification, extinction, and ecological change in this lineage.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 769–838.     

A Phylogenetic Analysis of Ilyocryptus Sars, 1862 (Cladocera: Ilyocryptidae)

Kotov and Štifter (2006) revised the taxonomy of the genus Ilyocryptus Sars, 1862 (Branchiopoda: Anomopoda: Ilyocryptidae) and concluded that 28 species can be recognized as valid for the world fauna. In order to test phylogenetic relationships between the species and to contribute to a better understanding of the genus, a cladistic (using branch‐and‐bound search) analysis was conducted for 25 Ilyocryptus species (two of them with two subspecies). In total, 32 morphological characters were used. An evolutionary‐morphological interpretation of the results was also made. Phylogenetic analysis revealed that Ilyocryptus elegans is an earlier derived member of the genus, while other species are separated into two main branches: agilis ‐ and sordidus ‐lines (without reference to type of moulting). Species with incomplete moulting and species with complete moulting do not form separate clusters, suggesting an independent origin of incomplete moulting within different species groups. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

Palaeoreas,Majoreas, and Stryfnotherium gen. nov. (Mammalia: Artiodactyla: Bovidae) from the Late Miocene of Greece

A new comparative study of the two crania, LGPUT RZO-03 and LGPUT RZO-68 attributed to Palaeoreas zouavei Bouvrain, 1980 from the Late Miocene of Axios Valley, Greece reveals great morphological and metric differences between them, which require the exclusion of LGPUT RZO-68 from this species and even from Palaeoreas Gaudry, 1861. This decision leaves unsupported many diagnostic features of P. zouavei applied by previous authors and along with recent data from Bulgaria put in doubt the validity of P. zouavei as a whole. As a result, the genus Palaeoreas is re-defined as monotypic and the cranial morphometric changes of P. lindermayeri (Wagner, 1848) are discussed on a temporal basis. An updated taxonomic analysis confirms the presence of P. lindermayeri in the Late Turolian mammal assemblage of Dytiko-1 (Axios Valley, Greece) but rejects its occurrence at Dytiko-2. Instead, a re-examination of old cranial specimens and a study of new ones suggest the presence in both sites of Majoreas Kostopoulos, 2004 a Late Miocene taxon previously mostly known from the eastern banks of the Aegean Sea. A reassessment of the LGPUT RZO-68 cranium, and its comparison with Palaeoreas and Criotherium Forsyth-Major, 1891, suggests that it deserves taxonomic distinction at genus and species levels: Stryfnotherium exophthalmon.     

Nouveau matériel de Leptobos (Smertiobos) crassus (Artiodactyla, Mammalia) du Pléistocène inférieur à Renzidong (Chine de l’Est)

New remains of Leptobos (Smertiobos) crassus have been identified among the fossils excavated from a new early Pleistocene site at the Renzidong in Anhui Province, Eastern China. It is a Leptobos bearing frontal appendices with a simple curvature and its distal part turns higher than the other species of the subgenus. The present study shows that the first appearance of Leptobos in China is in the zone equivalent to the European MNQ 16b, later than its first appearance in Western Europe (MNQ 16a); and the genus Leptobos can serve as index fossil of the lower Pleistocene in China (2.5-0.78 Ma, Chinese definition).     

牛科(哺乳纲:偶蹄目)动物与生境利用有关的适应形态模式

对广义牛科动物颅后骨骼的多元变量分析揭示了牛科生境利用和体型之间的骨学特征。利用逐步分辨分析方法和一个基于机器学习的决策树方法鉴别了每种生境中牛科动物颅后解剖结构的形态特征。从110个广义牛科动物测量了43个指标进行了这项分析。利用所有主要肢骨测量值和以单根肢骨测量为主的测量值获得的分辨函数和决策树可以完美地区分适应开阔生境、森林和山地的牛科动物(在所有分析中得到了100 %正确的再分类)。由于调整的函数仅涉及到很小的颅后骨骼测量集,这些函数可以应用于研究考古学和古生物学发掘物中保存的不完整标本。这些表征生境利用的生态适应函数与那些用颅齿部性状建立、用于推测牛科动物食物选择的函数结合,具有刻画已灭绝的分类类群的古个体生态学和重建古环境的潜力。我们还分析了多元回归是否较单一因子回归表现出较高的预测能力,并提出了从每一种单根主要肢骨测量的颅后形态变量得到的最好代数函数