Ameniscomys selenoides, an enigmatic aplodontiid rodent from the Early Miocene of Central Europe: a revision of the taxon based on new fossil evidence

Explanations of tropical intercontinental disjunctions involving South America and Africa typically invoke vicariance of western Gondwanan biotas or long-distance dispersal. However, many plant groups originated and diversified well after the last direct connection between Africa and South America (ca. 105 million years before the present [mybp]), and it is unlikely that long-distance dispersal accounts for the distribution of all of these. A less commonly invoked explanation, the boreotropics hypothesis, indicates that some tropical disjunctions arose during the Tertiary via high-latitude land connections when northern forests supported tropical vegetation. Malpighiaceae are widely distributed across Africa and South America and have been explained as ancient ‘‘Gondwanian aborigines’’ (i.e., vicariants of western Gondwanan biotas) or more recent ‘‘American colonists’’ (i.e., long-distance dispersalists from South America into the Old World). Fossil and phylogenetic evidence from clock-independent estimates of molecular divergence times indicate that Malpighiaceae originated in South America during the latest Cretaceous (ca. 68 mybp), in isolation from Africa, and that six amphi-Atlantic disjunctions within the family occurred during three major episodes: late Paleocene (ca. 60 mybp), latest Eocene‐earliest Oligocene (ca. 34‐31 mybp), and early Miocene (ca. 21‐17 mybp). These age estimates reject a Gondwanan origin for Malpighiaceae, and strict dispersal scenarios ignore paleoclimate, paleoland configurations, and fossil evidence that indicates that the family once inhabited northern latitudes. Instead, these data suggest that Paleocene‐Oligocene amphi-Atlantic disjunct groups in Malpighiaceae moved into North America from South America via the Caribbean Basin, crossed the North Atlantic into Eurasia, and subsequently reached the Old World Tropics during warm intervals when land configurations would have facilitated this migration. Whether Miocene migrations of evergreen thermophilic Malpighiaceae proceeded via northern latitudes or long-distance dispersal is less clear.

The concept of 'archaeophytes' (alien taxa which became established in a study area before AD 1500) is widely used in floristic analyses in central and northern Europe, but few authors have applied it to the British flora. Six criteria for the recognition of archaeophytes are outlined, drawing upon evidence of fossil and recent history, current habitat and European and extra-European distribution. These are used to identify 157 probable archaeophytes in Britain. Only five of these are known from fossil records from the Neolithic; most are first recorded in archaeological contexts in the Late Bronze Age, Iron Age, Roman or Medieval periods. As a group, archaeophytes (unlike neophytes) have declined in Britain in the 20th century. Comparison of the accepted status of these species in the Czech Republic, Germany, Poland and Finland demonstrates that over 50% are treated as archaeophytes in central Europe, but in Finland many are absent or only present as casual introductions. Species regarded as archaeophytes in these countries but as natives in Britain are also reviewed. The indirect nature of the evidence used to identify archaeophytes means that it is usually impossible to be certain about the history of a species; in particular, archaeophytes which have successfully invaded semi-natural habitats are likely to be overlooked as natives. The suggestion that a species is an archaeophyte is best regarded as a hypothesis to be tested by further studies. There is considerable scope for archaeological investigations aimed at addressing these botanical problems.

Galaxiid fishes are commonly assumed to exhibit a Gondwanan distribution, but little fossil evidence has been available to support or refute this assumption until now. Here we report on three species of fossil Galaxias, represented by exquisitely preserved, almost entire skeletons, together with jaw and skull fragments, from several Miocene lacustrine deposits in Otago, South Island, New Zealand. These are the only certain Galaxias fossils, and provide unequivocal evidence that the genus was abundant, and taxonomically and ecologically diverse in the New Zealand region before 23 million years ago. One new species, Galaxias effusus, from an Early Miocene diatomite deposit near Middlemarch is described on the basis of low vertebral count, short caudal peduncle, expansive dorsal, caudal and anal fins, rounded to truncate caudal fin, with 15 principal caudal fin rays. The intact skeletons of G. effusus confirm that they inhabited a deep, stratified maar lake with cool, anoxic bottom water. The maar lake was completely landlocked, from sedimentological evidence, and this species must have been non‐migratory. Disarticulated fish jaws and skull fragments from Early Miocene mudstones near Bannockburn are likely to be from a second large species of Galaxias on the basis of dentary, premaxilla and parasphenoid size and morphology. These disarticulated fish remains from a regionally extensive inland lake system indicate decomposition and/or predation in shallow nearshore environments, but we have no evidence yet of migratory behaviour. New specimens of G. kaikorai Whitley from Late Miocene diatomite deposits near Dunedin confirm its morphological similarity to the living G. brevipinnis. The occurrence and diversity of galaxiids in lakes of early and Late Miocene age in southern New Zealand can be explained by either oceanic dispersal across the Southern Hemisphere at any time during the Cenozoic, relict Gondwanan distributions or, most likely, an ancient Gondwanan origin overprinted by Cenozoic dispersal.

Patterns seen in other Australian flora have led to hypotheses that early Miocene shifts in climate drove rapid radiation of major taxonomic groups such as Eucalyptus. Little is known about absolute dates and rates for Australian monocots, particularly grasses. I tested this early Miocene radiation hypothesis for Australian grasses using a calibrated phylogeny of the endemic stipoid genus Austrostipa and an analysis of diversification rates. The phylogeny was developed from a Bayesian likelihood analysis of the nuclear internal transcribed spacers region, and three calibration points were set based on fossil evidence. The results indicate that the genus arose in the early Miocene and underwent a species radiation, but the rate of diversification was not rapid compared to the current rate or to those of other taxa. Following an 8 million year period of fast molecular evolution but no taxonomic radiation, diversification rates have been constant for the past 15 million years. Comparable measures such as the gamma statistic can be used across taxa to make general conclusions about evolutionary rate constancy.

Molecular phylogenetic studies that have included sirenians from the genera Trichechus, Dugong, and Hydrodamalis have resolved their interrelationships but have yielded divergence age estimates that are problematically discordant. The ages of these lineage splits have profound implications for how to interpret the sirenian fossil record—including clade membership, biogeographic patterns, and correlations with Earth history events. In an effort to address these issues, here we present a total evidence phylogenetic analysis of Sirenia that includes living and fossil species and applies Bayesian tip-dating methods to estimate their interrelationships and divergence times. In addition to extant sirenians, our dataset includes 56 fossil species from 106 dated localities and numerous afrotherian outgroup taxa. Genetic, morphological, temporal, and biogeographic data are assessed simultaneously to bring all available evidence to bear on sirenian phylogeny. The resulting time-tree is then used for Bayesian geocoordinates reconstruction analysis, which models ancestral geographic areas at splits throughout the phylogeny, thereby allowing us to infer the direction and timing of dispersals. Our results suggest that Pan-Sirenia arose in North Africa during the latest Paleocene and that the Eocene evolution of stem sirenians was primarily situated in the Tethyan realm. In the late Eocene, some lineages moved into more northern European latitudes, an area that became the source region for a key trans-Atlantic dispersal towards the Caribbean and northern-adjacent west Atlantic. This event led to the phylogenetic and biogeographic founding of crown Sirenia with the Dugongidae-Trichechidae split occurring at the Eocene-Oligocene boundary (~33.9 Ma), temporally coincident with the onset of dropping global sea levels and temperatures. This region became the nexus of sirenian diversification and supported taxonomically-rich dugongid communities until the earliest Pliocene. The Dugonginae-Hydrodamalinae split occurred near Florida during the early Miocene (~21.2 Ma) and was followed by a west-bound dispersal that gave rise to the Pacific hydrodamalines. The late middle Miocene (~12.2 Ma) split of Dugong from all other dugongines also occurred near Florida and our analyses suggest that the Indo-Pacific distribution of modern dugongs is the result of a trans-Pacific dispersal. From at least the early Miocene, trichechid evolution was based entirely in South America, presumably within the Pebas Wetlands System. We infer that the eventual establishment of Amazon drainage into the South Atlantic allowed the dispersal of Trichechus out of South America no earlier than the mid-Pliocene. Our analyses provide a new temporal and biogeographic framework for understanding major events in sirenian evolution and their possible relationships to oceanographic and climatic changes. These hypotheses can be further tested with the recovery and integration of new fossil evidence.

Fossil pollen and spores preserved in organic‐rich claystones near the base of the palaeochannel hosting the Tertiary Yandi channel iron deposits (CID) provide an in situ constraint on the age of this world‐class orebody. This assemblage also aids understanding of the palaeoenvironment leading to the deposition and preservation of the host Marillana Formation. The provisional Early Oligocene age of claystones links the genesis of the Yandi CID to the profound disruption of global climates and oceanography at the Eocene‐Oligocene transition (Terminal Eocene Event) and provides circumstantial evidence that ore formation was linked to the development of a warm south‐flowing proto‐Leeuwin Current along the Pilbara coast. It is likely that rainfall increased but remained strongly seasonal (monsoonal?) as global climates warmed during the Late Oligocene — Early Miocene. The same fossil evidence indicates that wood fragments, the iron‐oxide‐replaced remains of which are a significant component of cemented goethitic gravel making up CID, came from Casuarinaceae‐ and Myrtaceae‐dominated sclerophyll communities lining the channel banks. Increasingly dry conditions since the late Early Miocene Climatic Optimum are likely to have contributed to the preservation of CID at Yandi.

The fossil record documenting the evolution of hylobatids is extremely poor, so details of their phylogenetic and geographic origins and subsequent evolutionary history are obscure. Based on molecular clock estimates, hylobatids diverged from other hominoids during the early Miocene , at ~19 Ma, and crown hylobatids originated at ~8 Ma. The oldest fossil hylobatid is Yuanmoupithecus from the late Miocene of China, dating to ~7–9 Ma, which represents the primitive sister taxon of crown hylobatids. The molecular and paleontological evidence indicates that there was a ghost lineage for the initial 10 myrs of hylobatid evolutionary history, with no trace of a fossil record. Hylobatids presumably originated in Africa during the early Miocene, but the timing of their arrival in Asia and their early geographic distribution is unknown. Since there are no suitable fossil precursors for Yuanmoupithecus at older sites in China, it is likely that stem hylobatids migrated northwards from Southeast Asia during the late Miocene, but the Neogene fossil record from this region is poorly documented. Hylobatids occur at a number of Pleistocene archaeological and paleontological sites throughout southern China and Southeast Asia, but they tend to be relatively rare elements of the primate fauna. These are generally referable to extant lineages and species, except for Bunopithecus sericus from the early or middle Pleistocene of China. This contribution reviews what is known about the evolutionary history of the hylobatids based on the fossil evidence, but since there is much that we do not know and cannot deduce about the phylogeny of hylobatids from the incomplete fossil record, a fuller appreciation of the evolutionary history relies on what can also be learned from comparative anatomy and molecular data.

Amber fossils of sooty moulds

Molecular phylogenies and historical biogeography of a circumtropical group of gastropods (Genus: Nerita): Implications for regional diversity patterns in the marine tropics

The Cenozoic strata in the areas of Machino-machi, Najimi-mura and Yanaida-mura, in Fugeshi-gun, Ishikawa Prefecture, are classified, on the basis of field observations, into the following formations, from lower to upper : Konosuyama volcanics : Comprised of andesite, basalt, dacite, agglomerates of andesite, basalt and dacite, and diorite., Part of the diorite is younger than the volcanics., Although the distribution of the various rock types has been mapped their stratigraphic sequence is not known., ., ., ., (stratigraphic relation unceertain) ., ., ., Toktunari formation : Comprised of pumiceous tuff and brecciated tuff with lenticular layers of tuffaceous sandstone, tuffaceous mudstone and conglomerate., Lignite seams and fossil shells are found in a few places., These rocks are intercalated between flows of andesite, basalt and dacite., ., ., ., unconformity ., ., ., Higashi-Innai formation : Comprised of mudstone, siltstone, sandstone and conglomerate in which lignite is intercalated., Fossils of Mollusca and Foraminifera are abundant., ., ., ., Awagura formation and Iwakurayama rhyolite : The Awagura formation is comprised of finely stratified white or light green colored pumiceous tuff, brecciated tuff and sandy tuff., The Iwakurayama rhyolite eruption is responsible for the large quantities of tuff in the Awagura formation, and consequently, the eruption took place during deposition of the Awagura., ., ., ., Najimi formation : Comprised of tuffaceous mudstone and tuffaceous siltstone, and locally contains pumice layers and laminated tuffaceous sandstone layers., Molluscan shells and sponge spicules occur throughout the formation., In the northern part of the areas, Radiolaria and Globigerina, are common and Conchocele is the outstanding megafossil., In the southern part of the areas, Foraminifera are abundant and Globigerina is also found., The occurrence of Globigerina suggests a free connection with the open sea., ., ., ., unconformity ., ., ., Okawa terrace deposits : Comprised of sand, gravel and clay in which fossils are absent., From fossil evidence, the Higashi-Innai formation, which has yielded Operculina, Miogypsina, Vicarya and others, is considered to be early Miocene in age., The Higashi-Innai formation contains rocks of marine and brackish-water facies as evidenced by fossils and lignite which contains sulphur., Among the geological structures of the district are several faults, and folding has occurred., Faults and fold axes trend east-west., The geological age of the folding and faults is considered post-Miocene and pre-Okawa time.,

An exceptionally well‐preserved collection of Tertiary opisthobranch molluscs from the Aquitaine Basin, France, includes species of the order Notaspidea [Umbraculum sanctipaulensis sp. nov., Tylodina perversa (Gmelin), Spiricella unguiculus Rang and Des Moulins, Berthella aquitaniensis sp. nov., Berthella ateles sp. nov.], of the order Anaspidea [Akera cf. bullata Mu¨ller, Floribella corrugata (Cossmann), Floribella cossmanni sp. nov., Floribella rozieri sp. nov., Limondinia ornata gen. et sp. nov.] and of the order Sacoglossa [Volvatella faviae sp. nov.]. BerthellaaquitaniensisB. atelesV. faviae are the first fossil records of the families Volvatellidae and Pleurobranchidae. Floribella plicifera (Cossmann) and F.corrugata, originally assigned to the genus Philine, belong to the genus Floribella and constitute the oldest records of this genus. The fossil evidence indicates that in Umbraculum laudunensis and U. sanctipaulensis the shell probably covered most of the animal, whereas in the Recent U. umbraculum the shell only covers the central portion of the body. Tylodina perversa could be an old species that appeared during the early Miocene, more than 21 Ma. The Recent shells of Akera bullata are indistinguishable from fossils as old as the mid Eocene, but it may be biologically unrealistic to consider them to be the same species. The European species of Floribella evolved from the bullomorph shells of the early Eocene forms to the elongate shells of the early Miocene. The genus Volvatella is another example of marine tropical disjoint distributions and an excellent ecological indicator.

The present study aimed to assess the consequences of tectonic events and temperature regime on the diversification of Indo-West Pacific (IWP) turban shell species. Bayesian and parsimony methods were used to construct a phylogenetic hypothesis using sequence data from three genes for the turban shell genus Turbo and for a larger data set including representatives of all known genera in the subfamily Turbininae. Phylogenetic analyses indicate that all IWP Turbo species form a single clade approximately 68 Myr in age, predating the closure of the Tethys Sea and therefore predating the physical separation of the IWP from other biogeographical regions. All but one of the IWP subgenera tested in Turbo also predate the closure of the Tethys Sea. Fossil evidence for Marmarostoma, the largest subgenus, confirms that at least some Turbo lineages currently restricted to the IWP were previously more widespread. The combination of the phylogeny with the fossil evidence suggests that present day diversity in IWP Turbo is the result of the evolutionary persistence within the IWP of several, morphologically distinct lineages, some of which were more widespread in the Oligocene. Some IWP lineages show significant increases in diversification in the early Miocene, probably as a result of the increased availability of both shallow-water habitats due to tectonic plate movements and increased carbonate platforms in the central IWP resulting from coincident diversification of zooxanthellate corals. The IWP is therefore behaving as both a cradle of diversity (with new species originating in situ) and a museum of diversity (with lineages that predate its isolation also being maintained). Bayesian and parsimony analyses of the subfamily recovered five clades and mapping the temperature regime (tropical or temperate) of each species onto the molecular tree using parsimony suggested that temperate habitat is an ancestral character in at least four of the clades. This result was also supported by Bayesian reconstruction of ancestral states. Astralium (the fifth clade) may also prove to have a temperate origin, but this could not be determined with certainty given the available data. The origin of diversity in tropical regions is of particular interest because it has been suggested that the tropics are the source of many evolutionary novelties and have provided a species pool, from which temperate regions were populated. The present study suggests that Turbininae may be an exception to this rule. The tree shape also suggests that temperature has had an effect on speciation rates; temperate Turbininae are apparently evolving more slowly or suffering more extinction than their tropical sister clades, which show greater diversity.

The flora of the south-western tip of southern Africa, the Cape flora, with some 9000 species in an area of 90,000 km2 is much more speciose than can be expected from its area or latitude, and is comparable to that expected from the most diverse equatorial areas. The endemism of almost 70%, on the other hand, is comparable to that found on islands. This high endemism is accounted for by the ecological and geographical isolation of the Cape Floristic Region, but explanations for the high species richness are not so easily found. The high species richness is accentuated when its taxonomic distribution is investigated: almost half of the total species richness of the area is accounted for by 33 'Cape floral clades'. These are clades which may have initially diversified in the region, and of which at least half the species are still found in the Cape Floristic Region. Such a high contribution by a very small number of clades is typical of island floras, not of mainland floras. The start of the radiation of these clades has been dated by molecular clock techniques to between 18 million years ago (Mya) (Pelargonium) and 8 Mya (Phylica), but only six radiations have been dated to date. The fossil evidence for the dating of the radiation is shown to be largely speculative. The Cenozoic environmental history of southern Africa is reviewed in search of possible triggers for the radiations, climatic changes emerge as the most likely candidate. Due to a very poor fossil record, the climatic history has to be inferred from larger scale patterns, these suggest large-scale fluctuations between summer wet (Palaeocene, Early Miocene) and summer dry climates (Oligocene, Middle Miocene to present). The massive speciation in the Cape flora might be accounted for by the diverse limitations to gene flow (dissected landscapes, pollinator specialisation, long flowering times allowing much phenological specialisation), as well as a richly complex environment providing a diversity of selective forces (geographically variable climate, much altitude variation, different soil types, rocky terrain providing many micro-niches, and regular fires providing both intermediate disturbances, as well as different ways of surviving the fires). However, much of this is based on correlation, and there is a great need for (a) experimental testing of the proposed speciation mechanisms, (b) more molecular clock estimates of the age and pattern of the radiations, and (c) more fossil evidence bearing on the past climates.

Evolutionary radiation of an inbreeding haplodiploid beetle lineage (Curculionidae, Scolytinae)

Knowing the age of lineages is key to understanding their biogeographic history. We aimed to provide the best estimate of the age of Cichorieae and its subtribes based on available fossil evidence and DNA sequences and to interpret their biogeography in the light of Earth history. With more than 1,550 species, the chicory tribe (Cichorieae, Asteraceae) is distributed predominantly in the northern Hemisphere, with centres of distribution in the Mediterranean region, central Asia, and SW North America. Recently, a new phylogenetic hypothesis of Cichorieae based on ITS sequences has been established, shedding new light on phylogenetic relationships within the tribe, which had not been detected so far. Cichorieae possess echinolophate pollen grains, on the surface of which cavities (lacunae) are separated by ridges. These lacunae and ridges show patterns characteristic of certain groups within Cichorieae. Among the fossil record of echinolophate pollen, the Cichorium intybus-type is the most frequent and also the oldest type (22 to 28.4 million years old). By using an uncorrelated relaxed molecular clock approach, the Cichorieae phylogenetic tree was calibrated with this fossil find. According to the analysis, the tribe originated no later than Oligocene. The species-rich core group originated no later than Late Oligocene or Early Miocene and its subtribes diversified no later than Middle/Late Miocene or Early Pliocene—an eventful period of changing geological setting and climate in the Mediterranean region and Eurasia. The first dispersal from Eurasia to North America, which resulted in the radiation of genera and species in North America (subtribe Microseridinae), also occurred no later than Middle or Late Miocene, suggesting the Bering land bridge as the route of dispersal.