From Africa via Europe to South America: migrational route of a species‐rich genus of Neotropical lowland rain forest trees (Guatteria,Annonaceae)

Aim Several recent studies have suggested that a substantial portion of today’s plant diversity in the Neotropics has resulted from the dispersal of taxa into that region rather than by vicariance. In general, three routes have been documented for the dispersal of taxa onto the South American continent: (1) via the North Atlantic Land Bridge, (2) via the Bering Land Bridge, or (3) from Africa directly onto the continent. Here a species‐rich genus of Neotropical lowland rain forest trees (Guatteria, Annonaceae) is used as a model to investigate these three hypotheses. Location The Neotropics. Methods The phylogenetic relationships within the long‐branch clade of Annonaceae were reconstructed (using maximum parsimony, maximum likelihood and Bayesian inference) in order to gain insight in the phylogenetic position of Guatteria. Furthermore, Bayesian molecular dating and Bayesian dispersal–vicariance (Bayes‐DIVA) analyses were undertaken. Results Most of the relationships within the long‐branch clade of Annonaceae were reconstructed and had high support. However, the relationship between the Duguetia clade, the Xylopia–Artabotrys clade and Guatteria remained unclear. The stem node age estimate of Guatteria ranged between 49.2 and 51.3 Ma, whereas the crown node age estimate ranged between 11.4 and 17.8 Ma. For the ancestral area of Guatteria and its sister group, the area North America–Africa was reconstructed in 99% of 10,000 DIVA analyses, while South America–North America was found just 1% of the time. Main conclusions The estimated stem to crown node ages of Guatteria in combination with the Bayes‐DIVA analyses imply a scenario congruent with an African origin followed by dispersal across the North Atlantic Land Bridge in the early to middle Eocene and further dispersal into North and Central America (and ultimately South America) in the Miocene. The phylogenetically and morphologically isolated position of the genus is probably due to extinction of the North American and European stem lineages in the Tertiary.     

A scolopocryptopid centipede (Chilopoda: Scolopendromorpha) from Mexican amber: synchrotron microtomography and phylogenetic placement using a combined morphological and molecular data set

The first scolopocryptopid centipede known from the fossil record is a specimen of the subfamily Scolopocryptopinae in Miocene amber from Chiapas, southern Mexico. It is described here as Scolopocryptops simojovelensis sp. nov. , displaying a distinct combination of morphological characters compared to extant congeners. Anatomical details of the fossil specimen were acquired by non‐invasive 3D synchrotron microtomography using X‐ray phase contrast. The phylogenetic position of the new species is inferred based on a combination of morphological data with sequences for six genes (nuclear 18S and 28S rRNA, nuclear protein‐coding histone H3, and mitochondrial 12S rRNA, 16S rRNA, and protein‐coding cytochrome c oxidase subunit I) for extant Scolopendromorpha. The data set includes eight extant species of Scolopocryptops and Dinocryptops from North America, east Asia, and the Pacific, rooted with novel sequence data for other blind scolopendromorphs. The molecular and combined data sets, analysed in a parsimony/direct optimization framework, identified a stable pattern of two main clades within Scolopocryptopinae. North American and Asian species of Scolopocryptops are united as a clade supported by both morphological and molecular characters. Its sister group is a Neotropical clade in which the type species of Dinocryptops is nested within a paraphyletic assemblage of Scolopocryptops species; Dinocryptops is placed in synonymy with Scolopocryptops. The strength of support for the relationships of extant taxa from the molecular data allow the Chiapas fossil to be assigned with precision, despite ambiguity in the morphological data; the fossil is resolved as sister species to the extant Laurasian clade. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 768–786.     

Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time

The grasses (Poaceae) are the fifth most diverse family of angiosperms, including 800 genera and more than 10 000 species. Few phylogenetic studies have tried to investigate palaeo‐biogeographical and palaeo‐ecological scenarios that may have led to present‐day distribution and diversity of grasses at the family level. We produced a dated phylogenetic tree based on combined plastid DNA sequences and a comprehensive sample of Poaceae. Furthermore, we produced an additional tree using a supermatrix of morphological and molecular data that included all 800 grass genera so that ancestral biogeography and ecological habitats could be inferred. We used a likelihood‐based method, which allows the estimation of ancestral polymorphism in both biogeographical and ecological analyses for large data sets. The origin of Poaceae was retrieved as African and shade adapted. The crown node of the BEP + PACCMAD clade was dated at 57 Mya, in the early Eocene. Grasses dispersed to all continents by approximately 60 million years after their Gondwanan origin in the late Cretaceous. PACCMAD taxa adapted to open habitats as early as the late Eocene, a date consistent with recent phytolith fossil data for North America. C₄ photosynthesis first originated in Africa, at least for Chloridoideae in the Eocene at c. 30 Mya. The BEP clade members adapted to open habitats later than PACCMAD members; this was inferred to occur in Eurasia in the Oligocene. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 543–557.     

The long‐term history of dispersal among lizards in the early Eocene: new evidence from a microvertebrate assemblage in the Bighorn Basin of Wyoming,USA

Abstract: Early Eocene mammal faunas of North America were transformed by intercontinental dispersal at the Paleocene–Eocene boundary, but lizard faunas from the earliest Eocene of the same area were dominated by immigrants from within the continent. A new lizard assemblage from the middle early Eocene of Wyoming sheds light on the longer‐term history of dispersal in relation to climate change. The assemblage consists of three iguanid species (including two new species possibly closely related to living Anolis), Scincoideus, ‘Palaeoxantusia’, four anguids, two species of an undescribed new anguimorph clade, Provaranosaurus and a varanoid (cf. Saniwa). Most North American glyptosaurin glyptosaurines are now referred to Glyptosaurus, and Glyptosaurus hillsi is given a new diagnosis. Scincoideus is otherwise known only from the mid‐Paleocene of Belgium, and the specimens described here are the first to document intercontinental dispersal to North America among lizards in the early Eocene. Like in mammals, some immigrant lizard lineages first appearing in the Bighorn Basin in the earliest Eocene persisted in the area long after the Paleocene–Eocene thermal maximum, but other immigrants appear to have been restricted to the Paleocene–Eocene thermal maximum.     

Phylogenetics and taxonomy of the New World leafy spurges,Euphorbia section Tithymalus (Euphorbiaceae)

The 480 species of leafy spurges, Euphorbia subgenus Esula, represent the main temperate radiation in the large genus Euphorbia. This group is distributed primarily in temperate Eurasia, but with smaller, disjunct centres of diversity in the mountains of the Old World tropics, in temperate southern Africa and in the New World. The majority of New World diversity (32 species) occurs in a single section, section Tithymalus. We analysed sequences of the nrITS and plastid ndhF, trnH‐psbA, trnS‐trnG and trnD‐trnT regions to reconstruct the phylogeny of section Tithymalus and to examine the origins and diversification of the species native to the New World. Our results indicate that the New World species of section Tithymalus form a clade that is sister to the widespread, weedy E. peplus. The New World species fall into two primary groups: a ‘northern annual clade’ from eastern North America and a diverse clade of both annual and perennial species that is divided into three subgroups. Within the second group, there is a small ‘southern annual clade’ from Texas and northern Mexico, a perennial ‘Brachycera clade’ from the western United States and northern Mexico, and a perennial ‘Esuliformis clade’ from montane areas of Mexico, Guatemala, Honduras and the Caribbean island of Hispaniola. Ancestral state reconstructions indicate that the annual habit probably evolved in the ancestor of E. peplus and the New World clade, with a subsequent reversal to the perennial habit. In conjunction with this phylogenetic framework, the New World species of section Tithymalus are comprehensively reviewed. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 191–228.     

Phylogenetics and biogeography of the parasitic genus Thesium L. (Santalaceae), with an emphasis on the Cape of South Africa

Thesium is a large genus of parasitic shrubs belonging to tribe Thesieae of Santalaceae. It has a principally Old World distribution, with the greatest diversity being found in southern Africa. Little is known about the relationships within Thesium or its relationships with its closest relatives. In this article, we present a first estimate of species‐level phylogenetic relationships in Thesium based on internal transcribed spacer (ITS) and trnL–trnF sequence data, and use this to explore the biogeographical history of the group. One hundred and four samples representing 72 Thesium spp. were included in a phylogenetic analysis. Plastid and combined data resolve Thesium as paraphyletic relative to Thesidium and Austroamericium with high posterior probability and bootstrap support. ITS sequence data place Thesidium as sister to a large Thesium clade, but with weak support. Ancestral range reconstruction and dating analysis suggest a southern African origin for the group, with a crown age of 39.1 ± 11.9 Mya, followed by dispersal into Europe and South America. A large clade of Cape species split in the Miocene from a clade comprising tropical species (25.5 ± 7.3 Mya) with the diversification of extant species beginning at 16.7 ± 6.3 Mya. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 435–452.     

A reassessment of Eocene parrotlike fossils indicates a previously undetected radiation of zygodactyl stem group representatives of passerines (Passeriformes)

In the past years, various Eocene fossil birds were described as stem group representatives of the zygodactyl Psittaciformes (parrots). These birds show quite disparate morphologies, which cast some doubt on the correct assignment of all of them to the psittaciform stem group. A reassessment of their affinities is further needed, because it was recently proposed that among extant birds, Psittaciformes and Passeriformes (passerines) form a clade and that passerines possibly derived from a zygodactyl ancestor. Here, phylogenetic analyses are performed, which for the first time also include representatives of the Zygodactylidae, the extinct zygodactyl sister taxon of the Passeriformes. The early Eocene Psittacopes was originally described as a stem group representative of Psittaciformes. However, none of the present analyses supported psittaciform affinities for Psittacopes and instead recovered this taxon in a clade together with zygodactylids and passerines. Also part of this clade are the early Eocene taxa Pumiliornis and Morsoravis, and it is detailed that Psittacopes and the long‐beaked and presumably nectarivorous Pumiliornis, with which it has not yet been compared, are very similar in their postcranial osteology. The present analysis corroborates the hypothesis of a zygodactyl stem species of passerines. To account for these results, Psittacopes is here assigned to a new higher‐level taxon and a new name is also introduced for the clade including Zygodactylidae and Passeriformes.     

<Emphasis Type="Italic">Altingioxylon hainanensis</Emphasis> sp. nov.: earliest fossil wood record of the family Altingiaceae in Eastern Asia and its implications for historical biogeography

A new species, Altingioxylon hainanensis, is described from the Eocene Changchang Formation of the Changchang Basin on Hainan Island, South China. It is the first record of a fossil wood assigned to Altingiaceae found in China, and the most ancient evidence of wood for this family in eastern Asia. The new species is similar to A. rhodoleioides, known since the Miocene in India and Java Island, and to Altingia hisauchii from the Miocene to Pliocene of Japan. The close resemblance between these species and Liquidambar sp., known from the Middle Miocene of western North America, provides additional evidence for the migration of their ancestors from Asia to North America across the Bering land bridge during the Miocene. Distinctions in ray sizes between the eastern Asian specimens and their contemporaries from Europe to Kazakhstan is suggested as a result of the divergence between the large eastern Asian clade and the North American–west Asian clade within Altingiaceae during the Eocene–Oligocene. The presence of crystals in ray cells may be considered an ancestral condition that persists in the eastern Asian lineages up to the extant Altingia and Semiliquidambar, but which was lost in other Altingiaceae in the course of evolution.     

A new Transantarctic relationship: morphological evidence for a Rheidae–Dromaiidae–Casuariidae clade (Aves,Palaeognathae, Ratitae)

Although ratites have been studied in considerable detail, avian systematists have been unable to reach a consensus regarding their relationships. Morphological studies indicate a basal split separating Apterygidae from all other extant ratites, and a sister‐group relationship between Rheidae and Struthionidae. Molecular studies have provided evidence for the paraphyly of the Struthionidae and Rheidae, with respect to a clade of Australasian extant ratites. The position of the extinct Dinornithidae and Aepyornithidae also remains hotly debated. A novel pattern of diversification of ratites is presented herein. The phylogenetic analysis is based on 17 taxa and 129 morphological characters, including 77 new characters. The resultant tree yields a sister‐group relationship between New Zealand ratites (Apterygidae plus Dinornithidae) and all other ratites. Within this clade, the Aepyornithidae and Struthionidae are successive sister taxa to a new, strongly supported clade comprising the Rheidae, Dromaiidae, and Casuariidae. The link between South American and Australian biotas proposed here is congruent with numerous studies that have evidenced closely related taxa on opposite sides of the Southern Pacific. These repeated patterns of area relationships agree with current knowledge on Gondwana break‐up, which indicates that Australia and South America remained in contact across Antarctica until the earliest Tertiary. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 641–663.     

A new primitive alligatorine from the Eocene of Thailand: relevance of Asiatic members to the radiation of the group

Despite the report of fragmentary remains from China, the fossil record of primitive Alligatorinae is unbalanced with most of its members reported from North America and Europe. Here, we describe cranial, mandibular, and some postcranial elements of a new alligatorine taxon from the Late Eocene of Krabi Basin, southern Thailand. Krabisuchus siamogallicus gen. et sp. nov. was a small animal probably not surpassing 2 m in total length. Despite deformation, the remains show a dorsally elevated cranium similar to Arambourgia gaudryi and Osteolaemus tetraspis. The new alligatorine is characterized by a very short snout and a blunt dentition in the posterior region. Krabisuchus siamogallicus is the first fossil alligatorine from Asia to be incorporated into cladistic analysis. Results indicate a close relationship of the newtaxon to Procaimanoidea, Arambourgia gaudryi, and Allognathosuchus polyodon, all of them being sister groups to the Alligator clade. Alligatorine were widespread as early as the Late Eocene across the northern hemisphere. Scenarios of alligatorine dispersal during the Palaeogene are discussed in comparison with the European and North American alligatorine fossil record. Basal alligatorines were probably land‐dwelling animals contrary to modern‐day crocodylians. Such a mode of life may have allowed this group to colonize vast territories, especially during periods of global warm climates. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 608–628.     

A new kinosternoid from the Late Cretaceous Hell Creek Formation of North Dakota and Montana and the origin of the <Emphasis Type="Italic">Dermatemys mawii</Emphasis> lineage

A nearly complete turtle shell from the Late Cretaceous (Maastrichtian) Hell Creek Formation of Slope County, North Dakota, represents the most complete remains to date of a Mesozoic kinosternoid turtle and a new species, Hoplochelys clark nov. sp. The new taxon is diagnosable from other representatives of Hoplochelys by the plesiomorphic placement of the humeral/femoral sulcus behind the hyo/hypoplastral suture and the autapomorphic development of an interrupted median (neural) keel. All six previously named Paleocene (Puercan and Torrejonian) representatives of Hoplochelys lack diagnostic characters and are synonymized as Hoplochelys crassa. A phylogenetic analysis reveals that Hoplochelys spp. and Agomphus pectoralis are most parsimoniously placed within Kinosternoidea along the phylogenetic stem of the extant Mesoamerican River Turtle Dermatemys mawii, extending that taxon’s stem lineage from the early Eocene to the late Maastrichtian. The two primary crown lineages of Kinosternoidea are thus known from the Mesozoic and split prior to the late Campanian. The presence of a thickened cruciform plastron, true costiform processes, only three inframarginals, and the reduction of the medial contact of the abdominals are synapomorphies of Chelydroidea, the clade formed by Chelydridae and Kinosternoidae.     

New fossil mousebird (Aves: Coliiformes) with feather preservation provides insight into the ecological diversity of an Eocene North American avifauna

Coliiformes (mousebirds) are represented by just six extant species. These species, restricted to sub‐Saharan Africa, are all primarily frugivorous and are among the most sedentary of living birds. Previously described fossil Coliiformes preserving feather traces share the short, rounded wing shape of extant mousebirds. Along with osteological evidence, these observations have been proposed to support poor sustained flight capabilities across the stem mousebird lineage. We report a new species of Coliiformes from the early Eocene (51.66 ± 0.09 Ma) Fossil Butte Member of the Green River Formation, represented by one of the comparatively few fossils from these deposits preserving carbonized traces of the wing and tail feathering. Feather traces indicate an elongate, tapering wing shape similar to that of some extant aerial insectivores, and suggestive of a capacity for sustained and agile open‐air flight. Traces of the rectrices reveal the tail accounted for approximately two‐thirds of the total length of the bird, a proportion similar to that in living mousebirds. Phylogenetic analysis places the new species as a stem representative of Coliiformes, demonstrating for the first time that the two major clades of Coliiformes – Sandcoleidae and Colii – co‐occurred at Fossil Lake. Based on the recovered phylogeny, as well as the osteology and feathering of extant and fossil Coliiformes, the wing shape of the new species is interpreted as apomorphic. In addition to documenting unexpected morphological specialization within stem‐lineage Coliiformes, the new species adds yet another taxon to the emerging reconstruction of the diverse Paleogene avifauna from the tightly dated and nearly synchronous fossiliferous deposits of the Fossil Butte Member. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 685–706.     

Molecular phylogeny and biogeography of Astilbe (Saxifragaceae) in Asia and eastern North America

Phylogenetic analyses were conducted for Astilbe (Saxifragaceae), an Asian/eastern North American disjunct genus, using sequences of nuclear ribosomal internal transcribed spacer (ITS) and plastid matK, trnL‐trnF and psbA‐trnH regions. The monophyly of Astilbe is well supported by both ITS and plastid sequences. Topological incongruence was detected between the plastid and the ITS trees, particularly concerning the placement of the single North American species, A. biternata, which may be most probably explained by its origin involving hybridization and/or allopolyploidy with plastid capture. In Astilbe, all species with hermaphroditic flowers constitute a well‐supported clade; dioecious species form a basal grade to the hermaphroditic clade. Astilbe was estimated to have split with Saxifragopsis from western North America at 20.69 Ma (95% HPD: 12.14–30.22 Ma) in the early Miocene. This intercontinental disjunction between Astilbe and Saxifragopsis most likely occurred via the Bering land bridge. The major clade of Astilbe (all species of the genus excluding A. platyphylla) was inferred to have a continental Asian origin. At least three subsequent migrations or dispersals were hypothesized to explain the expansion of Astilbe into North America, Japan and tropical Asian islands. The intercontinental disjunct lineage in Astilbe invokes a hybridization event either in eastern Asia or in North America. This disjunction in Astilbe may be explained by a Beringian migration around 3.54 Ma (95% high posterior density: 1.29–6.18 Ma) in the late Tertiary, although long‐distance dispersal from eastern Asia to North America is also likely. The biogeographical connection between continental Asia, Taiwan, the Philippines and other tropical Asian islands in Astilbe provides evidence for the close floristic affinity between temperate or alpine south‐western China and tropical Asia. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●● , ●●–●●.     

New morphological evidence supports congruent phylogenies and Gondwana vicariance for palaeognathous birds

There has been little agreement on the phylogeny of palaeognathous birds, with major differences amongst and between results from morphological and molecular data. Two recently published phylogenies using nuclear and mitochondrial DNA have substantial agreement in overall topology, with the ostrich as sister group of all other extant palaeognaths and a kiwi‐emu‐cassowary clade. Here I report a morphological phylogeny based mainly on new characters from the tongue apparatus and cranial osteology, with a theoretical ancestor as outgroup. A new interpretation of the evolution of the avian palate is included. This phylogeny is very similar to these recent molecular results; this is the first report of such congruence, and offers a credible basis for understanding the evolution of this clade. This phylogeny is fully consistent with a Gondwana vicariance model of evolution. Dates attributed from known geological events place the first extant radiation (ostrich) in the mid‐Cretaceous, and offer a means of calibration of future molecular clock investigations. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 959–983.     

Molecular phylogenetics and biogeography of the eastern Asian–eastern North American disjunct Mitchella and its close relative Damnacanthus (Rubiaceae,Mitchelleae)

Mitchella is a small genus of the Rubiaceae with only two species. It is the only herbaceous semishrub of the family showing a disjunct distribution in eastern Asia and eastern North America, extending to Central America. Its phylogeny and biogeographical diversification remain poorly understood. In this study, we conducted phylogenetic and biogeographical analyses for Mitchella and its close relative Damnacanthus based on sequences of the nuclear internal transcribed spacer (ITS) and four plastid markers (rbcL, atpB‐rbcL, rps16 and trnL‐F). Mitchella is monophyletic, consisting of an eastern Asian M. undulata clade and a New World M. repens clade. Our results also support Michella as the closest relative to the eastern Asian Damnacanthus. The divergence time between the two intercontinental disjunct Mitchella species was dated to 7.73 Mya, with a 95% highest posterior density (HPD) of 3.14?12.53 Mya, using the Bayesian relaxed clock estimation. Ancestral area reconstructions suggest that the genus originated in eastern Asia. The semishrub Mitchella appears to have arisen from its woody ancestor in eastern Asia and then migrated to North America via the Bering land bridge in the late Miocene. © 2013 The Linnean Society of London     

Systematics,evolution, and biogeography of the Pliocene stem meline badger Ferinestrix (Carnivora: Mustelidae)

Ferinestrix vorax is an extinct mustelid carnivoran of enigmatic relationships, known from a partial mandible and femur collected from the 3.2‐ and 3.6‐Myr‐old deposits of Hagerman Fossil Beds National Monument, Idaho, USA. Here, we report Ferinestrix rapax sp. nov. based on 80 remains of skull and dentition from a 3.1–3.6‐Myr‐old deposit of Udunga, Transbaikal, Russia. We demonstrate that Ferinestrix is a stem genus of the badger subfamily Melinae. This genus is distinctly larger and more carnivorous than any other total‐clade meline. We show that Ferinestrix originated in Asia and immigrated to North America no later than at the early (Zanclean) to late (Piacenzian) Pliocene transition, and that the North American F. vorax and Asian F. rapax underwent parallel evolution toward increased carnivory. © 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167 , 208–226.