Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla,Hippopotamoidea)

We present new material of the selenodont anthracothere Hemimeryx blanfordi from the Oligocene deposits of the Bugti Hills (Balochistan, Pakistan), collected between 1999 and 2002. This is the first undisputed Oligocene occurrence of the species, previously known from the early Miocene of Pakistan. Investigation of the molar enamel microstructure reveals a surprising mono-zonal Schmelzmuster, already detected in some middle to Late miocene selenedont anthracotheres. We include this observation combined with a morphological revision of H. blanfordi and a cladistic assessment of the dental evidence, to propose a new phylogenetic hypothesis regarding Hemimeryx and its close relatives. We confirm the clade including advanced bothriodontines, which we erect to a tribe rank and name Merycopotamini. The South Asian origin of Merycopotamini is consistent with hypothesized subsequent dispersal events of Merycopotamini from Asia to Africa.     

A New Rhinoceros From The Lower Miocene Of The Bugti Hills, Baluchistan, Pakistan: The Earliest Elasmotheriine

The stratigraphical position of the Dera Bugti area (Baluchistan, Pakistan), which yielded the remains described herein, is discussed. Dental and postcranial material attributed to Bugtirhinus praecursor gen. et sp. nov., from the lower Miocene (MN 3) of the same area, is described. This species is the oldest and most primitive member of the elasmotheriine group within Rhinocerotidae (Mammalia). A new diagnosis is established for the Elasmotheriini. Comparison with the other lower and middle Miocene elasmotheriine taxa leads to the tree [Bugtirhinus praecursor [Caementodon oettingenae [other elasmotheriine taxa]]]. An upper Oligocene–lowermost Miocene Asiatic differentiation is suggested for the group, strengthened by a first appearance datum in the Dera Bugti area (lower part of the MN 3). The origin and dispersal of the elasmotheriines throughout Eurasia are discussed and correlated with major faunal exchanges. K ey words : Bugti Hills, Pakistan, Rhinocerotidae, Elasmotheriini, Miocene, biostratigraphy, palaeobiogeography.     

New Alicornops (Rhinocerotidae) remains from Lower and Middle Siwaliks,Pakistan

Unpublished rhinocerotid remains from the Lower and Middle Siwaliks of Pakistan are described in this paper and recognized as two species of Alicornops. Alicornops complanatum (Heissig) is identified in the Dhok Pathan Formation of the Middle Siwaliks and Alicornops laogouense Deng in the Kamlial Formation of the Lower Siwaliks. The Dhok Pathan Formation levels with A. complanatum are roughly correlated with the late Miocene-Pliocene European mammal zones MN10-15. In turn, levels with A. laogouense of the Kamlial Formation would correlate with the middle–late Aragonian (middle Miocene) European MN5. The recognition of the Chinese species A. laogouense in the Potwar Plateau represents the first discovery of this taxon in Pakistan and increases the geographical and stratigraphic distributions of this species, and adds to the rhinocerotid association from the Siwaliks. In turn, the presence of A. complanatum in the Siwaliks of Potwar Plateau also enlarges its geographic distribution in Pakistan, as it was previously known from the Bugti Hills of Balouchistan. The absence of Alicornops from the Siwaliks in the Chinji and Nagri formations (between late MN5 and MN9 zones) might be due to an inadequate fossil record, as other rhinocerotid species are known from Kamlial to Dhok Pathan formations. However, the two recorded species of Alicornops could also reveal two independent migration waves as supported by the appearance of other taxa in different formations. A summary of fossil Cenozoic rhinocerotids from different areas of Pakistan is also presented.     

Neogene paleobiogeography and East African paleoenvironments: contributions from the Tugen Hills rodents and lagomorphs.

A minimum of 28 genera of rodents and one genus of lagomorph were recovered from the Tugen Hills, Baringo District, Kenya, from localities dating from over 15.5 to about 4.4 Ma. The middle Miocene (sites dated between 15.8 and 15.3 Ma) rodent fauna recovered primarily from the Kipsaramon site complex, Muruyur Formation, includes a mixture of characteristically early Miocene taxa, and more derived forms. Composition of the African rodent fauna changes dramatically with the introduction of myocricetodontines, democricetodontines, and dendromurines, immigrants primarily from southern Asia. In the Tugen Hills, these taxa are first found in the Kabasero localities, Ngorora Formation, at sites dating from 12.5-12.33 Ma. A second major change in the African rodent fauna reflects the introduction of murines, immigrants from southern Asia. In the Tugen Hills murines are first encountered at Kapcheberek, Lukeino Formation, dated to 5.9-5.7 Ma. One rodent genus from the Lukeino Formation (Arvicanthis), and two from the Tabarin locality, Chemeron Formation (Heliosciurus, Paraxerus; 4.5-4.4 Ma), represent the earliest records of these extant African genera. A cricetomyine from the Ngorora Formation (12.5 Ma) is likely the earliest report of this exclusively African group. One of the earliest African records of porcupines (Hystricide) is from the Lukeino Formation. Lagomorphs are poorly represented, but include one of the earliest African occurrences of the family Leporidae from the Mpesida Beds (bracketed by dates of 7-6.2 Ma), and possibly a new genus of leporid from the Kapcheberek locality. Analysis of the Tugen Hills small mammals in association with other African records suggests several episodes of dispersal between Africa and Eurasia during the middle and late Miocene. Rodents from Kipsaramon are indicative of forests in conjunction with more open habitats. Those from the Kapcheberek locality are suggestive of a savanna habitat. The rodents from the Tabarin locality suggest a woodland environment.     

An osteology‐based appraisal of the phylogeny and evolution of kangaroos and wallabies (Macropodidae: Marsupialia)

Macropodids are the most diverse group of marsupial herbivores ever to have evolved. They have been the subject of more phylogenetic studies than any other marsupial family, yet relationships of several key clades remain uncertain. Two important problem areas have been the position of the merrnine (Lagostrophus fasciatus) and the phylogenetic proximity of tree‐kangaroos and rock‐wallabies. Our osteological analysis revealed strong support for a plesiomorphic clade ( Lagostrophinae subfam. nov. ) containing Lagostrophus and Troposodon, which is likely to have originated in the early Miocene. The extinct short‐faced kangaroos (Sthenurinae) emerged in the middle Miocene as the sister lineage to a clade containing all other living kangaroos and wallabies (Macropodinae). New Guinea forest wallabies ( Dorcopsini trib. nov. ) are the most plesiomorphic macropodines; the other two main lineages include tree‐kangaroos and rock‐wallabies (Dendrolagini), and ‘true’ kangaroos and wallabies (Macropodini). These phylogenetic outcomes are broadly consistent with the results of recent molecular studies, although conflicts remain over the relative positions of some macropodins (e.g. Setonix, Onychogalea, and Wallabia). Given the presence of derived dendrolagins and macropodins in early Pliocene localities, it is probable that most macropodine genera originated in the late Miocene. Key functional–adaptive trajectories within the craniodental and locomotory systems of the dominant macropodid lineages represent varying responses to the spread of drier, open habitats following the Miocene Climatic Optimum. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 954–987.     

The role of Asia in the origin and diversification of hystricognathous rodents

In the absence of a comprehensive pre‐Oligocene fossil record, the origin and early evolution of hystricognathous rodents have long been the subject of much uncertainty. Baluchimyinae (Rodentia) were initially interpreted as a subfamily of the ctenodactyloid Chappatimyidae (sciurognathous), a group considered to be endemic to the Indian subcontinent and to be closely related to hystricognathous rodents. A newly discovered early Oligocene hystricognathous rodent, Bugtimys zafarullahi gen. n. et sp. n., described herein, from the Bugti Hills (Balochistan, Pakistan) sheds new light on the higher level taxonomy of the previously described Baluchimyinae. As a contribution to the phylogenetic debates regarding the origin of Hystricognathi, we present a cladistic assessment of the dental evidence for the Palaeogene hystricognathous rodent cladogenesis. Our phylogenetic results consistently support the monophyly of the Hystricognathiformes clade (including Tsaganomyidae plus Hystricognathi) of which baluchimyine rodents are clearly members. There is, however, no support for the monophyly of a baluchimyine clade. Nonetheless, ‘baluchimyines’ are for the moment reinterpreted as Hystricognathi incertae sedis. Hystricognathous rodents appear to be well diversified at least since the early Oligocene, both in Africa and South America (phiomorphs and caviomorphs, respectively), and also now in south Asia. Furthermore, our phylogenetic results support close relationships between early hystricognathous and Asian ‘ctenodactyloid’ rodents, which clearly points to an Asian origin for Hystricognathi. In this phylogenetic framework, ‘baluchimyines’ and tsaganomyids are representatives of an initial phase of diversification of hystricognathous rodents in Asia. Oligocene phiomorphs and caviomorphs (sister groups) seem therefore to share a common ‘Asian’ hystricognathous ancestor. This reinforces the possibility that the early dispersal of hystricognathous rodents to South America was not from Africa but from Asia.     

Out of Africa? A dated molecular phylogeny of the cicada tribe Platypleurini Schmidt (Hemiptera: Cicadidae), with a focus on African genera and the genus Platypleura Amyot & Audinet‐Serville

The Platypleurini is a large group of charismatic cicadas distributed from Cape Agulhas in South Africa, through tropical Africa, Madagascar, India and eastern Asia to Japan, with generic diversity concentrated in equatorial and southern Africa. This distribution suggests the possibility of a Gondwanan origin and dispersal to eastern Asia from Africa or India. We used a four‐gene (three mitochondrial) molecular dataset, fossil calibrations and molecular clock information to explore the phylogenetic relationships of the platypleurine cicadas and the timing and geography of their diversification. The earliest splits in the tribe were found to separate forest genera in Madagascar and equatorial Africa from the main radiation, and all of the Asian/Indian species sampled formed a younger clade nested well within the African taxa. The tribe appears to have diversified during the Cenozoic, beginning c. 50–32 Ma, with most extant African lineages originating in the Miocene or later, well after the breakup of the Gondwanan landmass. Biogeographical analysis suggests an African origin for the tribe and a single dispersal event founding the Asian platypleurines, although additional taxon sampling and genetic data will be needed to confirm this pattern because key nodes in the tree are still weakly supported. Two Platypleurini genera from Madagascar (Pycna Amyot & Audinet‐Serville, Yanga Distant) are found to have originated by late Miocene dispersal of a single lineage from Africa. The genus Platypleura is recovered as polyphyletic, with Platypleura signifera Walker from South Africa and many Asian/Indian species apparently requiring assignment to different genera, and a new Platypleura concept is proposed with the synonymization of Azanicada Villet syn.n. The genera Orapa Distant and Hamza Distant, currently listed within separate tribes but suspected of platypleurine affinity, are nested deeply within the Platypleurini radiation. The tribe Orapini syn.n . is here synonymized while the tribe Hamzini is pending a decision of the ICZN to preserve nomenclatorial stability.     

Dioscoreaceae fossils from the late Oligocene and early Miocene of Ethiopia

Dioscorea section Lasiophyton leaflets from the late Oligocene (27.23 Ma) and Tacca leaves from the early Miocene (21.73 Ma) of north‐western Ethiopia greatly expand the known fossil record of Dioscoreaceae and represent the earliest and only known records of the Afro‐Asian trifoliate, palmately veined yams (Dioscorea) and bat flowers (Tacca). Both fossils occur in volcaniclastic and clastic sediments associated with a high water table, and the palaeofloral assemblages are indicative of tropical moist forest formations. These fossils provide insight into the evolutionary history of the family in Africa during the mid‐Cenozoic and provide well‐dated taxa that can assist in phylogenetic analyses and evolutionary divergence studies for Dioscoreales and Dioscoreaceae. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 17–28.     

Miocene snakes from northeastern Kazakhstan: new data on the evolution of snake assemblages in Siberia

Abstract

The Neogene snake fauna from the central and eastern regions of Eurasia is still largely unknown. This paper reports on a unique snake fauna from the late middle Miocene of the Baikadam and Malyi Kalkaman 1 and 2 localities, northeastern Kazakhstan, which represents the best-documented Miocene snake assemblage in Central Asia. Previous studies admitted that snake fauna could be homogeneous over a large part of Eurasia during the Miocene, with the late middle to early late Miocene assemblages similar to snake assemblages that inhabited Europe in the late early and early middle Miocene. This assumption is partially supported by the presence of Texasophis bohemiacus and Coluber cf. hungaricus, as well as vipers of the ‘V. aspis’ complex. However, the presence of taxa which are (1) probably not related to European representatives (‘Colubrinae’ A and B), (2) probably never occurred in Central and Western Europe and (3) are closely related to species recently inhabiting southern Siberia (Elaphe aff. dione, Gloydius sp.) indicates that faunal dissimilarity was relatively high within Eurasia during the late middle Miocene. This assumption is in accordance with studies of small mammal assemblages which show a decreasing homogenity in the Eurasia in the course of the middle Miocene.     

Detarieae sensu lato (Fabaceae) from the Late Oligocene (27.23 Ma) Guang River flora of north‐western Ethiopia

New species of caesalpinioid legumes, Cynometra sensu lato and Afzelia, are described from the Late Oligocene (27.23 Ma) Guang River flora in north‐western Ethiopia. Both taxa show leaf characteristics that are shared with extant species in the Guineo‐Congolian, Sudanian and/or Zambezian regions of Africa today. The presence of these two species in Ethiopia during the Palaeogene provides further evidence of the importance of the legume tribe Detarieae in northern and north‐eastern Africa throughout much of the Cenozoic, even although the clade is poorly represented in these regions today. The fossil record documents a significant palaeogeographical and evolutionary history of Detarieae in Africa, especially compared with that of Europe and Anatolia. Based on this evidence, it is unlikely that significant diversification of extant African Detarieae took place on the Eurasian landmass. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 44–54.     

New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean

Rowson, B., Tattersfield, P. & Symondson, W. O. C. (2010). Phylogeny and biogeography of tropical carnivorous land‐snails (Pulmonata: Streptaxoidea) with particular reference to East Africa and the Indian Ocean. —Zoologica Scripta, 40, 85–98. A phylogeny is presented for the speciose, near pan‐tropical, carnivorous achatinoid land‐snail superfamily Streptaxoidea inferred from DNA sequences (two nuclear and two mitochondrial regions) from 114 taxa from Africa, the Indian Ocean, Asia, South America and Europe. In all analyses, Streptaxidae are monophyletic, while the (two to six) previously recognised subfamilies are polyphyletic, as are several genus‐level taxa including the most speciose genus Gulella, necessitating major taxonomic review. The Asian Diapheridae are sister to Streptaxidae, which forms several well‐supported clades originating in a persistent basal polytomy. Divergence dating estimates, historical biogeography, and the fossil context suggest a Cretaceous origin of these families, but suggest Gondwanan vicariance predated most radiation. The basal polytomy dates to the Paleogene and may correspond to a rapid radiation in Africa. There is evidence for multiple Cenozoic dispersals followed by radiation, including at least two from Africa to South America, at least two from Africa to Asia and at least two from Africa to Madagascar, indicating Cenozoic turnover in tropical snail faunas. The endemic Seychelles and Mascarene streptaxid faunas each are composites of early Cenozoic lineages and more recent dispersals from Africa, with no direct evidence for an Asian origin as currently proposed. Peak streptaxid diversity in East Africa is explained by Neogene speciation among a large number of coexisting ancient lineages, a phenomenon most pronounced in the Eastern Arc‐Coastal Forests centre of endemism. This includes Miocene diversification in Gulella, a primarily East and South‐East African group which remains strikingly diverse even after unrelated ‘Gulella’ species are reclassified.     

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.     

Taxonomy,phylogeny, and diversity of the extinct Lesser Antillean rice rats (Sigmodontinae: Oryzomyini), with description of a new genus and species

Rice rats (Sigmodontinae: Oryzomyini) are abundant in the Late Quaternary fossil record and in Holocene pre‐Columbian archaeological middens across the Lesser Antilles. All of these rice rats are now extinct, and their regional diversity and systematics remain extremely poorly understood. We redescribe all of the region's rice rat taxa known from adequate diagnostic material (Megalomys desmarestii, Megalomys luciae, and Oligoryzomys victus), and describe a new genus and species, Pennatomys nivalis gen. et sp. nov. , from archaeological sites on St. Eustatius, St. Kitts, and Nevis, which formed a single larger island during Quaternary low sea‐level stands. Cladistic analysis supports the inclusion of O. victus within Oligoryzomys, and identifies Megalomys as a sister group of the large‐bodied genera Sigmodontomys or Sigmodontomys + Nectomys, suggesting that large body size in Megalomys represents phyletic gigantism rather than ‘island gigantism’. Megalomys and Pennatomys belong to an oryzomyine clade that has undergone remarkable radiation throughout the oceanic and continental‐shelf islands of the Neotropical region, but these genera do not represent a monophyletic group within the Nectomys subclade, indicating multiple over‐water colonization events of the Lesser Antillean island chain. Although Lesser Antillean rice rats were heavily exploited by prehistoric Amerindians, it is likely that most or all of these taxa survived until European arrival in the region. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 748–772.     

An assessment of the status of Polycirridae genera (Annelida: Terebelliformia) and evolutionary transformation series of characters within the family

A phylogenetic analysis was performed to determine the monophyly of non‐monotypic genera of the terebelliform family Polycirridae, i.e. Polycirrus, Amaeana, Lysilla, and Hauchiella, and the evolution of characters among members of this clade. The monotypic genera, Enoplobranchus and Biremis, were also included, together with members of both known species in Hauchiella. Representative species were included for remaining genera: 14 species of Polycirrus, six species of Amaeana, and six species of Lysilla. Out‐groups consisted of representatives of Spionidae, Cirratulidae, and Sabellariidae, as well as several species of Telothelepodidae. A total of 40 in‐ and out‐group species were coded for 50 subjects (‘characters’) and 117 subject–predicate relationships (‘states’). Although results are consistent with recent phylogenetic studies within Terebelliformia that suggest Polycirridae monophyly, only Hauchiella was found to be monophyletic, albeit part of the more inclusive clade comprising remaining polycirrid genera. Evolutionary transformation series are discussed for selected characters in relation to the non‐monophyly of Polycirrus, Lysilla, and Amaeana. Implications for the use of supraspecific taxa as ‘taxonomic surrogates’ are highlighted. The definition of Polycirridae is emended. © 2015 The Linnean Society of London     

Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia)

The Acochlidia are unique among opisthobranch gastropods in combining extremely high morphological and ecological diversity with modest species diversity. The phylogeny of acochlidians has never been addressed by cladistic means, as their evolution has remained unknown. This study gives a first overview on more than 150 biological and morphological characters that are potentially useful for phylogenetic analysis. Based on 107 characters, a parsimony analysis (PAUP) was performed for all 27 valid acochlidian species together with 11 (plus two) outgroup taxa. The resulting strict consensus tree shows a moderate overall resolution, with at least some bootstrap support for most resolved nodes. The Acochlidia are clearly monophyletic, and originate from an unresolved basal opisthobranch level. The Acochlidia split into the Hedylopsacea (Tantulum (Hedylopsis (Pseudunela (Strubellia (‘Acochlidium’, ‘Palliohedyle’))))) and Microhedylacea (Asperspina (Pontohedyle, ‘Parhedyle’, ‘Microhedyle’, (Ganitus, Paraganitus))). The formerly enigmatic Ganitidae, resembling sacoglossan opisthobranchs by having dagger‐like rachidian radular teeth, are likely to be highly derived microhedylids. The paraphyly of some of the traditionally recognized family level taxa induced a preliminary reclassification. From the phylogenetic hypothesis obtained, we conclude that the acochlidian ancestor was marine mesopsammic. The colonization of limnic systems occurred twice, independently: first in the Caribbean (with the development of the small interstitial Tantulum elegans), and second in the Indo‐Pacific, with a radiation of large‐sized benthic acochlidian species. The evolution of extraordinary reproductive features, such as hypodermic impregnation by a complex copulative aparatus in hedylopsaceans, cutaneous insemination via spermatophores in microhedylaceans, and gonochorism in Microhedylidae s.l. (including Ganitidae), is discussed. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 124–154.     

The Miocene floras of Iceland and their significance for late Cainozoic North Atlantic biogeography

A large number of plant macrofossils from several Middle to Upper Miocene localities from Iceland have been studied. The fossil material includes four ferns and fern allies, seven conifers, and about 40 species of flowering plants. Betula islandica and Salix gruberi are described as new species. Coniferous twigs previously ascribed to the genus Sequoia are shown to belong to Cryptomeria based on macro‐morphological and epidermal features. Fossil plants from Iceland are compared with coeval fossil taxa from Europe and North America and with living plants. The main finding is that the Miocene flora of Iceland belongs to a widespread Neogene northern hemispheric floral type including plants whose representatives are restricted to East Asia, North America and to western Eurasia at the present time. Previously inferred conspicuous similarities to North American modern equivalents appear to be misleading. The type of vegetation in four plant‐bearing sedimentary formations from the late Mid Miocene to Late Miocene, the 12 Ma Brjánslækur‐Seljá Formation, the 10 Ma Tröllatunga‐Gautshamar Formation, the 9–8 Ma Skarðsströnd‐Mókollsdalur Formation, and the 7–6 Ma Hreðavatn‐Stafholt Formation, corresponds to a humid temperate broadleaved (deciduous)–coniferous mixed forest dominated by Betulaceae, Fagaceae and Acer. Changes in species composition in the sedimentary formations reflect a shift from warm temperate (Cfa climate) to cool temperate (Cfb climate) conditions from the late Mid Miocene to the latest Miocene. This shift was connected to repeated phases of extinction and colonization. Specifically, one set of thermophilic taxa including Magnolia, Liriodendron, Sassafras and Comptonia went extinct between 12 and 10 Ma, and appears to have been replaced by another set of thermophilic taxa in the 10 Ma formation (Juglandaceae aff. Pterocarya/Cyclocarya, Rhododendron ponticum type). The 9–8 and 7–6 Ma formations are characterized by taxa that migrated to Iceland from Europe, such as Fagus gussonii, Betula cristata and Pterocarya fraxinifolia type. Although there is convincing evidence that plants colonized Iceland both from North America and Europe until 12 Ma, migration in the younger formations (9–8, 7–6 Ma) is suggested to have occurred mainly from Europe. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149 , 369–417.     

An early Miocene deep‐water decapod crustacean faunule from the Vienna Basin (Western Carpathians,Slovakia)

Abstract: A decapod crustacean faunule from the lower Miocene (upper Burdigalian, ‘Karpatian’) of the Slovakian part of the Vienna Basin comprise five new species: Callianopsis marianae (Ctenochelidae), Crosniera schweitzerae (Thomassiniidae), Agononida cerovensis and Munidopsis lieskovensis (both Galatheidae) plus Mursia harnicari (Calappidae). The new species of Callianopsis is the first undoubted member of the genus to be recorded from Europe; it is based on sexually dimorphic major and minor chelae as well as on portions of carapace and abdomen. Crosniera schweitzerae sp. nov. and Agononida cerovensis sp. nov. constitute the first fossil members of these genera. Additional material of an enigmatic crab, Styrioplax exiguus, and a re‐examination of the type material, confirms assignment of that genus to the subfamily Rhizopinae (family Pilumnidae). Palaeoecological data suggest that deposition of the levels (Lak?árska Nová Ves Formation) from which these taxa were collected took place under generally low‐energy, deep‐water conditions that were conducive to the preservation of delicate structures. Palaeobiogeographical affinities of the described taxa suggest a trans‐Atlantic migration during the early Miocene.     

The historical biogeography of Scabiosa (Dipsacaceae): implications for Old World plant disjunctions

Aim To reconstruct the temporal and biogeographical history of Old World disjunctions in Scabiosa (Dipsacaceae) and the timing of diversification in the Mediterranean Basin, in order to evaluate the importance of biogeographical and climatological history (particularly the onset of a mediterranean climate) in shaping Scabiosa distributions. Location Europe and the Mediterranean Basin, southern Africa and eastern Asia. Methods This study uses maximum‐likelihood and Bayesian phylogenetic analyses of chloroplast DNA (atpB–rbcL, trnL–trnF, trnS–trnG, psbA–trnH) and nuclear ribosomal DNA [internal transcribed spacer (ITS) and external transcribed spacer (ETS)] from 24 out of c. 37 ingroup taxa, beast molecular dating, and the dispersal–extinction–cladogenesis method (Lagrange ) to reconstruct ancestral geographical ranges and the timing of diversification of the major clades of Scabiosa. Results Biogeographical and divergence time reconstructions showed that Scabiosa originated during the Miocene and diversified in Europe, followed by independent movements into Asia and Africa. Several of the major clades were inferred to have radiated sometime between the late Miocene and early Pleistocene, a timeframe that encompasses the onset of the mediterranean climate in Europe. More recent middle–late Pleistocene radiations in the Mediterranean Basin and southern Africa have played a large role in Scabiosa diversification. Main conclusions Members of Scabiosa appear to have capitalized on adaptations to montane and/or dry conditions in order to colonize similar habitats in different biogeographical regions. The formation of the East African Rift mountains is potentially of great importance in explaining the southward migration of Scabiosa. The initial diversification of Scabiosa in Europe during the Miocene is not consistent with the initiation of the mediterranean climate, but may instead be associated with increased aridity and the retreat of subtropical lineages during this time. However, the radiation of some of the major subclades within Scabiosa may have been associated with an emerging mediterranean climate. More recent and rapid radiations in both the Mediterranean Basin and southern Africa highlight the probable importance of Pleistocene climate fluctuations in Scabiosa diversification.     

Cryptic species and Miocene diversification of Palaearctic naked‐toed geckos (Squamata: Gekkonidae) in the Indian dry zone

We sampled Palaearctic naked‐toed geckos from across their range in India and used two mitochondrial and two nuclear genes to reconstruct relationships within a global phylogeny. Published sequences of Peninsular Indian Hemidactylus allow us to contrast these two groups in dating analyses – providing insights into the history of the Indian dry zone. Palaearctic naked‐toed geckos first moved onto the Indian Plate in the Oligocene, with higher‐level diversification probably linked to collision of the Indian and Eurasian plates, and subsequent dispersal into‐India and diversification with increasing Miocene aridity. An independent gekkonid radiation with species in the dry zone, Hemidactylus diversified during the same period in Peninsular India. Our results demonstrate that dry zone taxa across India may date back to at least the Miocene, with a potential historical climatic barrier between the Indus and Peninsular Indian Divisions. ‘Cyrtopodion’ aravallense is revealed to be a complex with seven genetically and environmentally divergent lineages that began diversifying in the late Miocene, congruent with increased aridity in north‐western India. This discovery of cryptic diversity in the Indian dry zone represents the first terrestrial vertebrate radiation from north‐western central India and highlights how little we understand of the regions’ biodiversity, emphasizing the need for systematic geographic sampling and multiline evidence to reveal true patterns of diversity. The ancestor of ‘Cyrtopodion’ aravallense came into the nascent Indian dry zone in the Miocene and has since diversified, potentially in the absence of any sympatric scansorial rupicolous geckos. Cyrtopodion scabrum represents a unique case of commensalism and shows phylogeographic structure in its presumed native range. The taxonomic implications of our study include a number of undescribed species, recognition of ‘Cyrtopodion’ as a distinct lineage and the non‐monophyly of Altiphylax.