Finds of fragmentary bird skeletons in the Middle Miocene of the northern Caucasus

The first anatomically assembled skeletal remains of Neogene birds in Russia have been found. The head and a fragment of the vertebral column of a duck (Anatidae) and a hind limb of a perching bird (Passeriformes) from the Middle Miocene of the Krasnodar Region (Tsurevsky Formation) comprise the earliest known Miocene birds from European Russia. The skull of a very small duck (smaller than any extant species of Eurasian ducks) shows a combination of morphological characters characteristic of the extant species of Tadorna and Nettapus, and could belong to a representative of the fossil genus Mioquerquedula. This discovery supports a separate generic status for small-sized middle Miocene anatids from Eurasia, and suggests that they were more primitive than the extant Anatinae.     

Dabbling ducks (Aves: Anatidae) from the Middle Miocene of Mongolia

A new genus and two new species of ducks (Aves: Anatidae) from the Middle Miocene Sharga locality are described. Mioquerquedula minutissima gen. et sp. nov. is a very small duck. Anas velox Milne-Edwards, 1868 from the Middle Miocene of France is transferred to the genus Mioquerquedula. Aix praeclara sp. nov. described here is the oldest record of the modern genus Aix. A revision of the previously described small duck Anas soporata Kurochkin, 1976 shows that only the specimens from the Sharga locality should be referred to this species. The status of other small ducks from the Neogene of Europe and North America is discussed. The diversity of herbivorous and diving ducks in the Sharga locality indicates that Miocene Shargyn Govi Lake was rich in food resources.     

On the Middle Miocene avifauna of Maboko Island,Kenya

The Middle Miocene sediments of Maboko Island (Lake Victoria) in western Kenya yielded numerous avian bones, which remained, however, little studied. The significance of this material is shown by the recent identification of an opisthocomiform bird. In the present study, further avian remains from Maboko Island are described. Most of the specimens belong to aquatic or semi-aquatic groups, of which some are closely related to taxa known from Early and Middle Miocene European avifaunas, that is, Nectornis cormorants (N. africanus nov. sp.) and Laricola-like Laromorphae. The fossil material also includes Ciconiidae (cf. Ciconia), Pelecanidae, Phoenicopteridae (Leakeyornis aethiopicus), Musophagidae, and a species of Ardeidae, which closely resembles the taxon Pikaihao from the Early Miocene of New Zealand. Some avian remains from Maboko Island belong to higher-level taxa unknown from the Middle Miocene of Europe. The occurrence of a giant Jacanidae (?Nupharanassa mabokoensis nov. sp.) is of particular interest, because these are globally absent in extant avifaunas and were previously only known from the Late Eocene/Early Oligocene of Egypt. Further unknown from contemporaneous European sites are small representatives of Jacanidae, Bucerotidae, and Alcedinidae, with the fossils of the latter two taxa being among the earliest published records of their respective groups. Several of the taxa that are common in contemporaneous European avifaunas have not been found in Maboko, and in spite of less pronounced climatic differences, Middle Miocene Afrotropical avifaunas already appear to have been distinct from contemporaneous European ones.     

Change in diversity,ecological significance and biogeographical relationships of the Mediterranean Miocene toothed whale fauna

The main evolutionary trend in the Mediterranean Miocene toothed whale fauna is related (1) to the change in diversity and (2) to the turnover in community structure. Diversity increases from Upper Aquitanian–Lower Burdigalian to Burdigalian–Langhian, when it reaches its maximum. Starting from this time, diversity decreases progressively. The Early Miocene (Upper Aquitanian–Lower Burdigalian) Mediterranean toothed whale fauna, as well as the extramediterranean ones, is characterised by a high number of endemic taxa and by the prevalence of longirostral forms living in estuarine-neritic environments. A more diversified fauna spreading in neritic and pelagic environments characterises the Burdigalian–Langhian age, while an increase in pelagic forms and the nearly complete disappearance of some archaic longirostral taxa is typical of the Serravallian–Messinian fauna. Decrease in diversity and disappearance of archaic longirostral taxa are also recorded, at more general scale, in the Late Miocene extramediterranean fossil bearing deposits. These events can be related to the progressive global climatic deterioration, starting from Middle Miocene. From a biogeographic point a view, we can outline some relationships between the Mediterranean and western North Atlantic Miocene faunas. Closer affinities are observed between the Baltringen fauna and the northern Atlantic one, because of the presence of the genera Pomatodelphis and Zarhachis (platanistids) in both areas. In the Miocene Mediterranean and in North Atlantic, the delphinids are apparently absent as well as other extant delphinoid groups even if erroneously recorded in the past.     

First fossils of Zygogynum from the Middle Miocene of Central Yunnan,Southwest China,and their palaeobiogeographic significance

The collision of Gondwana and Laurasia effected plant exchanges between the two continents and thus affected the evolution of the flora of Yunnan, Southwest China. Consequently, the plant diversity in Yunnan has been enriched such that it has attracted the attention of botanical researchers. In this paper, we describe winteraceous fossil leaves collected from the Middle Miocene in Central Yunnan. Compared with the leaves of today’s plants, they share cuticular features and leaf morphological characters with Zygogynum Baillon (Winteraceae); the fossil plant is designated as a new species, Zygogynum poratus n. sp. Liang and Zhou. Based on its historical and recent distribution, the genus probably had migrated into Yunnan from Gondwana during or before the Middle Miocene, spreading beyond its current geographic range.     

Middle Miocene Cephalotaxus (Taxaceae) from Yunnan,Southwest China,and its implications to taxonomy and evolution of the genus

The infrageneric taxonomy of the genus Cephalotaxus Siebold et Zuccarini ex Endlicher (Taxaceae) is controversial as morphological characters have been doubted to change coherently among species. Although the epidermal features are considered as important characters in the taxonomy of conifers, they have not been systematically studied in this genus. Neogene foliage shoots with well-preserved epidermal structure not only can provide systematic links to the extant species, but also have implications for the taxonomical definition of the extant species by their epidermal features. In this paper, the morphologies of leafy shoots combining epidermal structure of a Neogene Cephalotaxus, which was collected from the middle Miocene of Southwest China, and ten extant species (variants) were studied. Morphology as well as epidermal structures was compared among species (variants) in Cephalotaxus. Based on the comparisons, the taxonomical definition of extant species in genus Cephalotaxus are discussed. Our results indicate that leaf morphological characters in combination with the features of epidermal structure can distinguish species in genus Cephalotaxus. The fossil foliage shoots from the middle Miocene of Southwest China were assigned to a new species, C. maguanensis n. sp., which shows close affinities to the extant C. hainanensis. On the basis of leaf and epidermal information, we suggest there are seven extant species in Cephalotaxus. With global cooling and the uplift of the Qinghai-Tibet Plateau since the middle Miocene, the cooling climate induced the differentiation and southward expansions of C. maguanensis, and its extant offspring, C. hainanensis, is distributed to Hainan Island and the Indo-Chinese peninsula.     

The first known fossil record of pipehorses (Teleostei: Syngnathidae: Haliichthyinae) from the Miocene Coprolitic Horizon from the Tunjice Hills,Slovenia

The first known fossil specimens of pipehorses (Haliichthyinae) were unearthed from the Middle Miocene (Sarmatian) beds of the Coprolitic Horizon in the Tunjice Hills, Slovenia. These fossil pipehorses belong to a new genus and species Hippohaliichthys edis, which was similar to the extant species Haliichthys taeniophorus. The body morphology indicates that the described fossil pipehorses were also closely related to the pygmy pipehorse Hippotropiscis frenki and the seahorse Hippocampus sarmaticus, two taxa which were also found in the Coprolitic Horizon. The described fossil material of pipehorses indicates that seahorses evolved from a group of pipehorses that were similar in size and shape to extant and fossil pipehorses of the Haliichthyinae subfamily.     

Systematics and evolution of the Pan‐Alcidae (Aves,Charadriiformes)

Puffins, auks and their allies in the wing‐propelled diving seabird clade Pan‐Alcidae (Charadriiformes) have been proposed to be key pelagic indicators of faunal shifts in Northern Hemisphere oceans. However, most previous phylogenetic analyses of the clade have focused only on the 23 extant alcid species. Here we undertake a combined phylogenetic analysis of all previously published molecular sequence data (～ 12 kb) and morphological data (n = 353 characters) with dense species level sampling that also includes 28 extinct taxa. We present a new estimate of the patterns of diversification in the clade based on divergence time estimates that include a previously vetted set of twelve fossil calibrations. The resultant time trees are also used in the evaluation of previously hypothesized paleoclimatic drivers of pan‐alcid evolution. Our divergence dating results estimate the split of Alcidae from its sister taxon Stercorariidae during the late Eocene (～ 35 Ma), an evolutionary hypothesis for clade origination that agrees with the fossil record and that does not require the inference of extensive ghost lineages. The extant dovekie Alle alle is identified as the sole extant member of a clade including four extinct Miocene species. Furthermore, whereas an Uria + Alle clade has been previously recovered from molecular analyses, the extinct diversity of closely related Miocepphus species yields morphological support for this clade. Our results suggest that extant alcid diversity is a function of Miocene diversification and differential extinction at the Pliocene–Pleistocene boundary. The relative timing of the Middle Miocene climatic optimum and the Pliocene–Pleistocene climatic transition and major diversification and extinction events in Pan‐Alcidae, respectively, are consistent with a potential link between major paleoclimatic events and pan‐alcid cladogenesis.     

Fossil wood of <Emphasis Type="Italic">Laricioxylon erkovetskiense</Emphasis> sp. nov. (Pinaceae) from the Miocene of the Erkovetskii Brown Coal Field,Amur Region,Russia

A new species, Laricioxylon erkovetskiense sp. nov. (Pinaceae) identified on the basis of anatomical features of fossil wood from the Sazanka Formation (upper Middle Miocene–Upper Miocene) of the Erkovetskii Brown Coal Field (Amur Region) is described. The new species shares some characters with the extant larches Larix cajanderi and L. olgensis.     

Early Miocene catarrhine dietary behaviour: the influence of the Red Queen Effect on incisor shape and curvature

The early Miocene catarrhine fossil record of East Africa represents a diverse and extensive adaptive radiation. It is well accepted that these taxa encompass a dietary range similar to extant hominoids, in addition to some potentially novel dietary behaviour. There have been numerous attempts to infer diet for these taxa from patterns of dental allometry and incisor and molar microwear, however, morphometric analyses until now have been restricted to the post-canine dentition. It has already been demonstrated that given the key functional role of the incisors in pre-processing food items prior to mastication, there is a positive correlation between diet and incisal curvature (Deane, A.S., Kremer, E.P., Begun, D.R., 2005. A new approach to quantifying anatomical curvatures using High Resolution Polynomial Curve Fitting (HR-PCF). Am. J. Phys. Anthropol. 128(3), 630-638.; Deane, A.S., 2007. Inferring dietary behaviour for Miocene hominoids: A high-resolution morphometric approach to incisal crown curvature. Ph.D. Dissertation. The University of Toronto.). This study seeks to re-examine existing dietary hypotheses for large-bodied early Miocene fossil catarrhines by contrasting the incisal curvature for these taxa with comparative models derived from prior studies of the correlation between extant hominoid incisor curvature and feeding behaviour. Incisor curvature was quantified for 78 fossil incisors representing seven genera, and the results confirm that early Miocene fossil catarrhines represent a dietary continuum ranging from more folivorous (i.e., Rangwapithecus) to more frugivorous (i.e., Proconsul) diets, as well as novel dietary behaviours that are potentially similar to extant ceboids (i.e., Afropithecus). Additionally, early Miocene fossil catarrhine incisors are less curved than extant hominoid incisors, indicating a general pattern of increasing mesio-distal and labial curvature through time. This pattern of morphological shifting is consistent with the Red Queen Effect (Van Valen, L., 1973. A new evolutionary law. Evol. Theory 1, 1-30), which predicts that taxa that are removed from one another by geological time, although potentially having similar diets, may exhibit differing degrees of a similar dietary adaptation (i.e., differing degrees of incisal curvature).     

Mid-Tertiary elapid snakes (Squamata, Colubroidea) from Riversleigh, northern Australia: early steps in a continent-wide adaptive radiation

Vertebral and cranial remains of elapid snakes have been collected from fossil assemblages at Riversleigh, north-west Queensland, Australia; most are Miocene but one may be late Oligocene and another as young as Pliocene. The oldest specimen (probably the oldest elapid yet known anywhere) is a vertebra that can be referred provisionally to the extant taxon Laticauda (Hydrophiinae, sensu Slowinski and Keogh, 2000), implying that the basal divergences among Australasian hydrophiine lineages had occurred by the early Miocene, in contrast to most previous estimates for the age of this geographically isolated adaptive radiation. Associated vertebrae and jaw elements from a Late Miocene deposit are described as Incongruelaps iteratus nov. gen. et sp., which has a unique combination of unusual derived characters otherwise found separately in several extant hydrophiine taxa that are only distantly related. Associated vertebrae from other sites, and two parietals from a possibly Pliocene deposit, suggest the presence of several other taxa distinct from extant forms, but the amount of material (and knowledge of variation in extant taxa) is currently insufficient to diagnose these forms. The Tertiary elapids of Riversleigh thus appear to be relatively diverse taxonomically, but low in abundance and, with one exception, not referable to extant taxa below the level of Hydrophiinae. This implies that the present diversity of hydrophiine elapids (31 recognized terrestrial genera, and approximately 16 marine) represents the result of substantial extinction as well as the “cone of increasing diversity” that could be inferred from phylogenetic studies on extant forms.     

Descriptions and phylogenetic relationships of a new genus and two new species of Oligo‐Miocene cormorants (Aves: Phalacrocoracidae) from Australia

Tertiary cormorant fossils (Aves: Phalacrocoracidae) from Late Oligocene deposits in Australia are described. They derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations in the Lake Eyre and Lake Frome Basins, South Australia, respectively. A new genus, Nambashag gen. nov. , with two new species ( Nambashag billerooensis sp. nov. , 30 specimens; Nambashag microglaucus sp. nov. , 14 specimens), has been established. Phylogenetic analyses based on 113 morphological and two integumentary characters indicated that Nambashag is the sister taxon to the Early Miocene Nectornis miocaenus of Europe and all extant phalacrocoracids. As Nambashag, Nectornis, and extant phalacrocoracids constitute a strongly supported clade sister to Anhinga species, the fossil taxa have been referred to Phalacrocoracidae. Sulids and Fregata were successive sister taxa to the Phalacrocoracoidea, i.e. phalacrocoracids + Anhinga. As phalacrocoracids lived in both Europe and Australia during the Late Oligocene and no older phalacrocoracid taxa are known, the biogeographical origin of cormorants remains unanswered. The phylogenetic relationships of extant taxa were not wholly resolved, but contrary to previous morphological analyses, considerable concordance was found with relationships recovered by recent molecular analyses. Microcarbo is sister to all other extant phalacrocoracids, and all Leucocarbo species form a well‐supported clade. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 277–314.     

Khoratpithecus piriyai, a Late Miocene hominoid of Thailand

A Khoratpithecus piriyai lower jaw corresponds to a well-preserved Late Miocene hominoid fossil from northeastern Thailand. Its morphology and internal structure, using a microcomputed tomography scan, are described and compared to those of other known Miocene hominoids. It originated from fluviatile sand and gravel deposits of a large river, and was associated with many fossil tree trunks, wood fragments, and large vertebrate remains. A biochronological analysis by using associated mammal fauna gives an estimated geological age between 9-6 Ma. The flora indicates the occurrence of a riverine tropical forest and wide areas of grassland. K. piriyai displays many original characters, such as the great breadth of its anterior dentition, suggesting large incisors, large lower M3, a canine with a flat lingual wall, and symphysis structure. Several of its morphological derived characters are shared with the orangutan, indicating sister-group relationship with that extant ape. This relationship is additionally strongly supported by the absence of anterior digastric muscle scars. These shared derived characters are not present in Sivapithecus, Ankarapithecus, and Lufengpithecus, which are therefore considered more distant relatives to the orangutan than Khoratpithecus. The Middle Miocene K. chiangmuanensis is older, displays more primitive dental characters, and shares several dental characters with the Late Miocene form. It is therefore interpreted as its probable ancestor. But its less enlarged M3 and more wrinkled enamel may suggest an even closer phylogenetic position to orangutan ancestors, which cannot yet be supported because of the incomplete fossil record. Thus Khoratpithecus represents a new lineage of Southeast Asian hominoids, closely related to extant great ape ancestors.     

Enters the shrew,some considerations on the Miocene palaeobiogeography of Iberian insectivores

The fossil record of the Iberian insectivores forms a subset of those found in central Europe. Comparison of the late early to early late Miocene record of the two areas shows that, particularly during the late Early Miocene, central European taxa have transient occurrences in Spain. Most taxa appear earlier and survive longer in central Europe. A notable exception is the gymnure Galerix, which extirpates earlier in central Europe, except for a transient occurrence in Germany just prior to its extinction. The main period of insectivore migrations is the late middle Miocene, although some of the taxa that enter remain restricted to the coastal areas. Overall, the pattern of distribution in time and space is best explained by the preference of insectivores for humid environments, as were found during the early Miocene and re-appeared at the end of the middle Miocene.     

Fossil Wood of <Emphasis Type="Italic">Ulmus priamurica</Emphasis> sp. nov. (Ulmaceae) from the Miocene of the Erkovetskii Brown Coal Field,Amur Region,Russia

A new species, Ulmus priamurica sp. nov. (Ulmaceae), from the deposits of the Sazanka Formation (upper Middle–Upper Miocene) of the Erkovetskii Brown Coal Field (Amur Region, Russia) is described based on anatomical features of fossil wood. The new species shows some wood anatomical characters of the extant elms U. japonica and U. americana. Fossil wood of Ulmus was found in the Amur Region for the first time.     

福建漳浦中中新世几种藓类植物研究*

亚洲新生代藓类植物化石非常稀少,漳浦琥珀是目前已知的我国低纬度地区唯一含苔藓植物化石的新生代琥珀矿床。漳浦琥珀源自中中新世气候适宜期(~14-17 Ma)的龙脑香树脂,温暖潮湿的热带季雨林气候和广泛分布的龙脑香科植物,为藓类植物的生长、多样性演化和三维立体保存提供了条件。本研究共报道了漳浦县中中新世藓类植物化石3个属种,包括:白发藓属Leucobryum Hampe、牛舌藓属Anomodon Hook.et Taylor和扁枝藓属Homalia(Brid.)B.S.G.。首次利用三维X射线断层扫描技术(Micro-CT)研究了白发藓属化石,发现其与现生种短枝白发藓Leucobryum humillimum Cardot非常相似,因此将其命名为短枝白发藓(相似种)Leucobryum cf.humillimum。此外,当前牛舌藓属Anomodon和扁枝藓属Homalia化石是东亚地区的首例记录。漳浦琥珀中的藓类植物化石揭示了中中新世时期苔藓植物的多样性,丰富了东亚低纬度地区新生代苔藓植物的化石记录。     

New carnivoran remains from the Late Miocene of Turkey

A great number of carnivoran remains from various late Miocene localities of Turkey, housed in the Natural History Museum of Aegean University, Bornova-Izmir, are described and compared with those from Eurasian localities for their determination. Thirteen different taxa were determined and some of them are recognized for the first time in Turkey or in their localities. Noteworthy is the presence of Dinocrocuta gigantea, which was recognized for the first time in Turkey, found in the early Vallesian locality Bayraktepe II, and the presence of the otter Sivaonyx hessicus which is a rare taxon in Eurasia, known from Eppelsheim, Germany and from Küçükçekmece and Bayraktepe II, Turkey. The hyaenid Adcrocuta cf. eximia was recognized for the first time in the Vallesian of Middle Sinap. The new determined carnivoran taxa provided some biochronological information which helps to date the localities or confirm their age.     

A contribution to the herpetofauna from the late Miocene of Gritsev (Ukraine)

Excavations in the locality Gritsev (karstic fillings of the upper Miocene age in a limestone quarry in Shepetovsky District, Khmelnitsky Region, Ukraine) revealed, since 1983 when excavations started, a rich vertebrate fauna that includes also amphibians and lizards. The material is rich enough to assess individual and developmental variation within some taxa (e.g., Pseudopus pannonicus). Although the fragmentary nature of the disarticulated bones does not sometimes allow precise taxonomic assignment, morphological differences in, e.g., dental characteristics suggest the presence of some other ones, not formally named forms. Thus, the list of the amphibian and reptilian taxa from Gritsev is by far not complete, but their osteological characteristics are representative to such a degree that they can provide information for comparisons with other European Miocene herpetofaunas.     

Middle Miocene elasmotheriine Rhinocerotidae from China and Mongolia: taxonomic revision and phylogenetic relationships

Eight elasmotheriine rhinocerotid species have been described from the Middle Miocene of China and Mongolia. In this paper a revised taxonomy is presented based on direct observation and comparison of the available material. A phylogenetic analysis based on 282 morphological characters has led to the reappraisal of Procoelodonta Matthew, 1931, Caementodon Heissig, 1972 and Huaqingtherium Huang & Yan, 1983. The genus Procoelodonta is split into three subgenera: P.  ( Procoelodonta ) Matthew, 1931, P.  ( Begertherium ) Beliaeva, 1971, and P.  ( Pasalarhinus subg. n). The genus Caementodon is split into two subgenera: C.  ( Caementodon ) Heissig, 1972 and C.  ( Beliajevina ) Heissig, 1974. Four species are assumed to have occurred in the Middle Miocene within the area studied: Procoelodonta ( Procoelodonta ) mongoliense (Osborn, 1924), ' P. ' ( Begertherium ) borissiaki (Beliaeva, 1971), ' Caementodon ' ( Beliajevina ) fangxianense (Yan, 1979) and Huaqingtherium lintungense (Zhai, 1978) (=' Caementodon tongxinensis ' Guan, 1988 =' Huaqingtherium qiui ' Guan, 1993 =' Hispanotherium tungurense ' Cerdeño, 1996). Shennongtherium hypsodontus Huang & Yan, 1983 is removed from the Elasmotheriina, owing to dental characters which suggest that it is a teleoceratine. The distribution of the main characters later seen in Elasmotherium is briefly discussed. The persistence and diversity of the Elasmotheriina throughout the Middle Miocene help explain how minute brachyodont animals gave rise to the mammoth-sized hypsodont Elasmotherium.     

The parrots (Aves: Psittaciformes) from the Middle Miocene of Sansan (Gers,Southern France)

The occurrence of Archaeopsittacus sp. (Psittaciformes) in the fossil deposits of Sansan (France) is reported, testifying to a survival of this genus in the Middle Miocene, as Archaeopsittacus verreauxi was described from the Early Miocene of Saint-Gérand-le-Puy (France) and was recorded only from its type locality. The data discussed here indicate the presence of two parrot species from Sansan, as it is the type locality of another parrot species, Pararallus dispar, only known from this locality. The differences between the humeri of these two taxa are described in detail, together with the differences from the other European fossil parrot species. The presence of more than one species of parrot in the same locality is not rare, but in Europe it is recorded in Sansan for the second time. Evidence for parrots also confirms the paleoenvironmental reconstruction of Sansan, as parrots are primarily arboreal species. Archaeopsittacus also represents one of the few common elements between the Early and Middle Miocene European bird assemblages.