Molecular phylogeny and historical biogeography of the large carpenter bees, genus Xylocopa (Hymenoptera: Apidae)

The biogeographical history of major groups of bees with worldwide distributions have often been explained through hypotheses based on Gondwanan vicariance or long distance dispersal events, but until recently these hypotheses have been very difficult, if not impossible, to distinguish. New fossil data, comprehensive information on Mesozoic and Cenozoic coastline positions and the availability of phylogenetically informative DNA markers now makes it feasible to test these hypotheses for some groups of bees. This paper presents historical biogeographical analyses of the genus Xylocopa Latreille, based on phylogenetic analyses of species belonging to 22 subgenera using molecular data from two nuclear genes, elongation factor‐1α (EF‐1α) and phosphoenolpyruvate carboxykinase (PEPCK), combined with previously published morphological and mitochondrial data sets. Phylogenetic analyses based on parsimony and likelihood approaches resulted in several groups of subgenera supported by high bootstrap values (>85%): an American group with the Oriental/Palaearctic subgenera Nyctomelitta and Proxylocopa as sister taxa; a geographically diverse group (Xylocopa s.l); and a group consisting of African and Oriental subgenera. The relationships among these three clades and the subgenus Perixylocopa remained unresolved. The Oriental subgenus Biluna was found to be the sister group of all other carpenter bee subgenera included in this study. Using a relaxed molecular clock calibrated using fossil carpenter bees, we show that the major splits in the carpenter bee phylogeny occurred well after the final breakup of Gondwanaland (the separation of South America and Africa, 100 Mya), but before important Miocene fusion events. Ancestral area analysis showed that the genus Xylocopa most likely had an Oriental‐Palaearctic origin and that the present world distribution of Xylocopa subgenera resulted mainly from independent dispersal events. The influence of Pleistocene glaciations on carpenter bee distributions is also discussed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 249–266.     

Globally basal centres of endemism: the Tasman-Coral Sea region (south-west Pacific), Latin America and Madagascar/South Africa

The New Zealand wrens (Acanthisittidae) are basal in passerine birds and in New Caledonia, the closest country to New Zealand, Amborella is basal in angiosperms. A review of molecular studies indicates that 29 other locally or regionally endemic clades around the Tasman and Coral Sea basins have cosmopolitan or globally widespread sister groups. Other areas that have local endemics basal to diverse global groups include Borneo, Madagascar/South Africa/Tanzania, southern China–Taiwan–Japan, and different parts of Latin America, especially the Guayana Plateau and western Mexico. Basal clades are widely interpreted as ancestral and their location is generally taken to represent a centre of origin for the group as a whole. In the present study, basal groups are treated simply as small (less speciose) sister groups. The Tasman and western Mexico/Caribbean regions have important biogeographic and tectonic ties with each other and with the central Pacific. Large igneous provinces (Ontong Java, Hikurangi‐Manihiki, and Gorgona Plateaus) formed in the central Pacific in the Cretaceous. Fossil wood is found on the Ontong Java Plateau, and formerly emergent seamounts up to 24 km across occur on Hikurangi Plateau. Many terranes in New Zealand, New Caledonia, New Guinea and western America represent former island arcs (or their products) that formed in the central Pacific and later accreted to the Pacific margins. Long‐term survival of taxa as metapopulations on the ephemeral volcanic islands and atolls of plate margins and fissures may explain the biogeographical connections among the Tasman region, the central Pacific, and the accreted terranes of western America. Branching sequences in cladograms and phylogenetic trees have been interpreted as dispersal events, but instead probably indicate the sequence of differentiation in an already widespread ancestor. Major disjunctions of tens of thousands of kilometres often occur between taxa at consecutive nodes on a tree and dispersal by physical movement is unlikely. The break between the globally basal centres and the rest of the world marks the initial site of differentiation of a widespread ancestor, with subsequent or more or less simultaneous differentiation occurring in other areas. Differentiation of the modern angiosperms, passerines and other groups first took place around the Tasman region, or at least the terranes now accumulated there, and then around other centres, especially Madagascar–South Africa and Mexico–north‐west South America. Angiosperms are now recognized as basal to extant gymnosperms and major tectonic dynamism around the globally basal centres during the Mesozoic, involving terrane accretion, orogeny, and rifting could have been involved with the last important modernization of angiosperms, birds and other groups. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96 , 222–245.     

Molecular phylogeny of Tornier's cat snake (Crotaphopeltis tornieri), endemic to East African mountain forests: biogeography, vicariance events and problematic species boundaries

Five isolated populations of the East African snake species Crotaphopeltis tornieri were investigated using mitochondrial sequence data from the genes cytochrome B and NADH2. The congeneric species Crotaphopeltis hotamboeia and three species of the closely related genera Dipsadoboa were used as outgroups. The results were analysed using unweighted parsimony and the reconstructed phylogeny revealed four well-defined evolutionarily significant units. Very high levels of genetic diversity was observed. The highest levels of divergence was observed between specimens from Mount Rungwe and the remaining populations. A similar pattern but with much lower divergence levels has been observed for the bird species Andropadus masukuensis . Together they indicate that Mt Rungwe is an important area for future conservation efforts. In the analysis of the combined data set C. tornieri appear as non-monophyletic with C. hotamboeia being the sister group to all C. tornieri populations except those on Mt Rungwe. The validity of C. tornieri and C. hotamboeia as species was thereby questioned by the data. However, the support for this is very weak and no nomenclatural consequences are suggested at present.     

Molecular phylogeny and historical biogeography of the genus Tuber, the 'true truffles'

Aim Various data sets and methods of analysis were combined to produce the first comprehensive molecular phylogeny of the genus Tuber and to analyse its biogeography. Location Europe, North Africa, China, Asia, North America. Methods Phylogenetic relationships among Tuber species were reconstructed based on a data set of internal‐transcribed spacer (ITS) sequences and various phylogenetic inference methods, specifically maximum parsimony, Bayesian analysis and neighbour joining. Tajima’s relative rate test showed that Tuber 18S rRNA, 5.8S rRNA, 5.8S‐ITS2 rRNA and β‐tubulin sequences evolved in a clock‐like manner. These genes, combined or not, were employed for molecular clock estimates after construction of linearized trees using mega 3.1. We reconstructed ancestral areas in the Northern Hemisphere by means of a dispersal–vicariance analysis (diva 1.1) based on current distribution patterns of the genus Tuber determined from the literature. Results The resulting molecular phylogeny divided the genus Tuber into five distinct clades, in agreement with our previously published studies. The Puberulum, Melanosporum and Rufum groups were diversified in terms of species and geographical distribution. In contrast, the Aestivum and Excavatum groups were less diversified and were located only in Europe or North Africa. Using a global molecular clock analysis, we estimated the divergence times for the origin of the genus and for the origin of several groups. diva inferred nine dispersal events and suggested that the ancestor of Tuber was originally present in Europe or was widespread in Eurasia. Equally optimal distributions were obtained for several nodes, suggesting different possible biogeographical patterns. Main conclusions Our analyses identified several discrepancies with the classical taxonomy of the genus, and we propose a new phylogenetic classification. According to molecular clocks, the radiation of the genus Tuber could have started between 271 and 140 Ma. Used in combination with the results obtained from time divergence estimates, this allows us to propose two equally probable scenarios of intra‐ and inter‐continental diversification of the genus according to the geographic distribution of the most recent common ancestor in Europe or Eurasia. The biogeographical patterns imply intra‐continental dispersal events between Europe and Asia and inter‐continental dispersal events between North America and Europe or Asia, which are compatible with land connections during the Tertiary.     

Minute moss beetles in the Southern Hemisphere: Molecular phylogeny,historical biogeography and habitat shifts (Coleoptera: Hydraenidae)

Minute moss beetles (Hydraenidae) are one of the most speciose and widespread families of aquatic Coleoptera, with an estimated 4000 extant species, found in the majority of aquatic habitats from coastal rock pools to mountain streams and from the Arctic Circle to the Antarctic islands. Molecular phylogenetic works have improved our understanding of the evolutionary history of the megadiverse Hydraena, Limnebius and Ochthebius in recent years, but most genera in the family have not yet been included in any phylogenetic analyses, particularly most of those which are restricted to the Southern Hemisphere. Using a multimarker molecular matrix, sampling over 40% of described species richness and 75% of currently recognized genera, we infer a comprehensive molecular phylogeny of these predominantly Gondwanan Hydraenidae. Whilst the genera we focus on are morphologically diverse, and currently classified across all four hydraenid subfamilies, our phylogenetic analyses suggest that these Gondwanan genera may instead constitute a single clade. As a result of our findings, the African genus Oomtelecopon Perkins syn.n. is shown to nest within Coelometopon Janssens, the New Zealand Homalaena Ordish syn.n. and Podaena Ordish syn.n. are synonymised with Orchymontia Broun, and the South African Pterosthetops Perkins syn.n. is synonymised with Prosthetops Waterhouse, resulting in Pterosthetopini Perkins syn.n. being synonymised with Prosthetopini Perkins. Mesoceratops Bilton & Jäch gen.n. is erected to accommodate six former members of Mesoceration Janssens, which is shown to be polyphyletic. We propose the replacement name Orchymontia ordishi Jäch & Bilton nom.n. for Homalaena dilatata Ordish, 1984 (now a junior homonym); altogether 39 new combinations are proposed. Our Bayesian divergence times infer an origin for this ‘Gondwana group’ of genera in Africa plus Madagascar in the mid-Cretaceous and suggest that both vicariant and dispersal processes, together with extinctions, have shaped the biogeographic history of these beetles in the Southern Hemisphere during the Cretaceous, resulting in geographically conserved extant lineages. Finally, we reconstruct ancestral habitat shifts across our phylogeny, revealing numerous changes in habitat occupancy in these genera, including multiple origins of fully terrestrial, humicolous taxa in different regions.     

Molecular phylogeny of the large carpenter bees, genus Xylocopa (Hymenoptera: apidae), based on mitochondrial DNA sequences

Carpenter bees, genus Xylocopa Latreille, a group of bees found on all continents, are of particular interest to behavioral ecologists because of their utility for studies of the evolution of mating strategies and sociality. This paper presents phylogenetic analyses based on sequences of two mitochondrial genes cytochrome oxidase 1 and cytochrome b for 22 subgenera of Xylocopa. Maximum-parsimony and maximum-likelihood methods were used to infer phylogenetic relationships. The analyses resulted in three resolved clades of subgenera: a South American group (including the subgenera Stenoxylocopa, Megaxylocopa, and Neoxylocopa), a group including the subgenera Xylocopa s.s. and Ctenoxylocopa, and an Ethiopean group (including the subgenera Afroxylocopa, Mesotrichia, Alloxylocopa, Platynopoda, Hoploxylocopa, and Koptortosoma). The relationships between the 11 other subgenera and the resolved clades are unclear. Within the Ethiopian group we found a clear separation of the African and the Oriental taxa and apparent polyphyly of the subgenus Koptortosoma. Using an evolutionary rate for ants, we investigated whether Gondwana vicariance or more recent dispersal events could best explain the present-day distribution of subgenera. Although some taxa show divergences that approach Gondwanan breakup times, most divergences between geographic groups are too recent to support a vicariance hypothesis.     

When Thailand was an island – the phylogeny and biogeography of mite harvestmen (Opiliones,Cyphophthalmi, Stylocellidae) in Southeast Asia

Aim To develop a comprehensive explanation for the biological diversity of Southeast Asia, especially in the Wallacea and Sundaland regions. This study focuses on a group of arachnids, mite harvestmen, which are thought to be an extremely old group of endemic animals that have been present in the region since most of its land supposedly formed part of the northern rim of the supercontinent Gondwana. Location Eastern Himalayas, Thai‐Malay Peninsula, Sumatra, Borneo, Java, Sulawesi, and New Guinea. Methods  Approximately 5.6 kb of sequence data were obtained from 110 South‐east Asian Cyphophthalmi specimens. Phylogenetic analyses were conducted under a variety of methods and analytical parameters, and the optimal tree was dated using calibration points derived from fossil data. Event based and paralogy‐free subtree biogeographical analyses were conducted. Results The Southeast Asian family Stylocellidae was recovered as monophyletic, arising on what is now the Thai‐Malay Peninsula and diversifying into three main clades. One clade (Meghalaya, here formally placed in Stylocellidae) expanded north as far as the eastern Himalayas, a second clade entered Borneo and later expanded back across the Sundaland Peninsula to Sumatra, and a third clade expanded out of Borneo into the entire lower part of Sundaland. Molecular dating suggested that Stylocellidae separated from other Cyphophthalmi 295 Ma and began diversifying 258 Ma, and the lineage that inhabits mostly Borneo today began diversifying between 175 and 150 Ma. Main conclusions The topology and molecular dating of our phylogenetic hypothesis suggest that Stylocellidae originated on Gondwana, arrived in Southeast Asia via the Cimmerian palaeocontinent, and subsequently diversified north, then south. Their present distribution in the Indo‐Malay Archipelago is explained largely by a diversification over the Sundaland Peninsula before western Sulawesi departed and the peninsula was extensively inundated.     

South Pacific biogeography,tectonic calibration,and pre‐drift tectonics: cladogenesis in Abrotanella (Asteraceae)

Abrotanella is the basal genus in the large tribe Senecioneae (Asteraceae) and has a disjunct distribution in Australasia and South America. A recent molecular phylogeny of the genus was used to investigate whether the main biogeographical patterns in the group could be related to the region's tectonic history in a coherent way. The phylogenetic/biogeographical breaks and overlaps in the genus imply a series of vicariance and range expansion events. Each of these can be related to one of the main tectonic events in the region, including assembly of the New Zealand terranes, crustal extension, and magmatism in Gondwana that preceded seafloor spreading, opening of the Tasman and Pacific basins, and transcurrent movement on the New Zealand Alpine fault. The coincident sequence indicates that pre‐drift tectonics and magmatism have been more important for the origin of trans‐Tasman and trans‐Pacific groups than the final rifting of Gondwana that led to their disjunction. For example, during the pre‐drift phase of break‐up, the Whitsunday volcanic province of Australia and the Median Batholith of New Zealand formed a large, active igneous belt. Its distribution is aligned with the break between New Zealand–south‐eastern Australia clades, and New Zealand–New Guinea clades. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

On the phylogeny of the Arminidae (Gastropoda, Opisthobranchia, Nudibranchia) with considerations of biogeography

In the past, the different genera of the family Arminidae have been diagnosed (mostly) on the basis of plesiomorphic characters and therefore their monophyly is questionable. The Arminidae are characterized by the autapomorphies 'possession of marginal sacs' and 'rhinophores with longitudinal lamellae'. The genus Heterodoris is excluded from the family. Two possible hypotheses about the phylogenetic relationships within the Arminidae are presented.
The Arminidae probably originated in the western Pacific, near Japan and have distributed in a westerly direction. The most primitive species ( Dermatobranchus ) are restricted mainly to the western Pacific region while the more derived species ( Armina ) have a world-wide distribution.     

Systematics and biogeography of the widespread Neotropical gekkonid genus Thecadactylus (Squamata), with the description of a new cryptic species

The systematics of the widespread neotropical gekkonid genus Thecadactylus is investigated using a combination of morphological, morphometric and mtDNA (cytochrome b ) sequence data, forming a total evidence dataset. The analysis tackles the common yet complex problem of a widespread taxon consisting of one or more cryptic species that are difficult to diagnose using morphology alone. The data are analysed using both maximum parsimony and Bayesian inference. Virtually all analyses resolve a well-supported south-western Amazonian clade distinct from the remainder of the recognized T. rapicauda . The south-western Amazonian clade is not only robustly supported, but also exclusive, geographically coherent and sufficiently distinct to warrant specific recognition. The new species is diagnosable on the basis of molecular sequences that are 23.0–26.9% divergent from those of T. rapicauda , and morphological evidence. Bayesian inference analysis robustly resolves meaningful and repeatable patterns of relationship. The biogeography of Thecadactylus is interpreted in the context of its two constituent species, and difficulties of resolving systematic and biogeographical patterns in widespread, cosmopolitan taxa are discussed. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 339–370.     

Phylogeny of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae) based on morphological and molecular evidence (CO1 and 16S), with a hypothesized diversification in the Pleistocene

The genus Ixchela Huber is composed of 20 species distributed from north‐eastern Mexico to Central America, including the five new species described here from Mexico: I xchela azteca sp. nov. , I xchela jalisco sp. nov. , I xchela mendozai sp. nov. , I xchela purepecha sp. nov. and I xchela tlayuda sp. nov. We test the monophyly and investigate the phylogenetic relationships among species of the genus Ixchela using morphological and molecular data. Parsimony (PA) analysis of 24 taxa and 40 morphological characters with equal and implied weights supported the monophyly of Ixchela with eight morphological synapomorphies. The PA analyses with equal and implied weights, and separate Bayesian inference (BI) analyses for the CO1 gene (506 characters), concatenated gene fragments CO1 + 16S (885 characters), morphology + CO1 (546 characters) and the combined evidence data set (morphology + CO1 + 16S) (925 characters) support the monophyly of Ixchela. Our preferred topology shows two large clades; clade 1 has a natural distribution in the Mesoamerican biotic component, whereas clade 2 predominates in the Mexican Montane biotic component. The genus Ixchela diverged in the late Miocene, and the divergence between the internal clades in the genus occurred in the late Pliocene; by contrast, most of the speciation events seem to have occurred mainly during the Pleistocene, where climatic changes brought on by repeated glaciations played an important role in the diversification of the genus. © 2015 The Linnean Society of London     

Historical biogeography of the catfish genus Hypostomus (Siluriformes: Loricariidae), with implications on the diversification of Neotropical ichthyofauna

Tropical South America possesses the largest ichthyofauna of any continental region. To test whether palaeohydrological changes may have been the causes of such diversification, the 'hydrogeological' hypothesis, the phylogenetic relationships of 51 representatives of the catfish genus Hypostomus (Siluriformes: Loricariidae) were inferred using mitochondrial D-loop haplotype sequences. Specimens were collected in all main tropical South American rivers systems east to the Andes. The major interrelationships found with the D-loop data were confirmed with a subset of 21 species using complete internal transcribed spacer (ITS) region sequences. The phylogenetic analysis indicate that the genus Hypostomus can be divided into four monophyletic clades. The historical biogeographical analysis of each of these clades allows the identification of seven major cladogenetic events. Using calibrated D-loop and ITS molecular clocks, date estimations were attributed to each of these cladogenetic events allowing a linkage between four of them with documented hydrogeological changes. Comparisons with published distribution patterns of unrelated fish groups indicate that several of the reconstructed and dated hydrogeological-cladogenetic events may have acted at a large scale on the diversification of Neotropical freshwater fish fauna during late Tertiary.     

Systematic revision,molecular phylogeny and biogeography of the antlion tribe Acanthoplectrini (Neuroptera: Myrmeleontidae: Dendroleontinae), with emphasis on the Oriental lineage

The tribe Acanthoplectrini (Myrmeleontidae: Dendroleontinae) includes a group of antlion genera widely distributed across the Australasian and Oriental regions. The intergeneric and interspecific relationships between or within the Australian and Oriental lineages of this tribe as well as their historical biogeography remain largely unexplored. Here, we present a molecular phylogenetic and biogeographic analyses of Acanthoplectrini to infer the diversification history of this tribe, with emphasis on the Oriental lineage. Both the Oriental and Australian lineages are monophyletic and recovered as sister groups. Ancestral area reconstruction suggests that the ancestor of Acanthoplectrini might have been once widely distributed from Indochina to Australia and then split into the Oriental and Australian lineages during the early-Miocene. Our analyses recovered northeastern Indochina and south China as the ancestral range of the Oriental Acanthoplectrini. During the mid-Miocene to the mid-Pliocene, orographic events such as the rising of mountain ranges (including the Himalayas) and the formation of major islands in southeastern Asia triggered several dispersal and vicariance events in the Oriental Acanthoplectrini, driving their speciation. We revise the classification of the Oriental Acanthoplectrini, establishing the new genus Paralayahima gen. n. , which is recovered sister to Layahima Navás. Moreover, we describe four new species of Layahima, Layahima aspoeckorum sp. n. , Layahima monba sp. n. , Layahima lhoba sp. n. and Layahima xinliae sp. n. , and we reinstate two previously synonymized species, Layahima melanocoris (Yang) stat. rev. and comb. n. and Layahima nebulosa Navás stat. rev.     

Molecular phylogeny and biogeography of the fern genus Pteris (Pteridaceae)

Background and Aims

Pteris (Pteridaceae), comprising over 250 species, had been thought to be a monophyletic genus until the three monotypic genera Neurocallis, Ochropteris and Platyzoma were included. However, the relationships between the type species of the genus Pteris, P. longifolia, and other species are still unknown. Furthermore, several infrageneric morphological classifications have been proposed, but are debated. To date, no worldwide phylogenetic hypothesis has been proposed for the genus, and no comprehensive biogeographical history of Pteris, crucial to understanding its cosmopolitan distribution, has been presented.

Methods

A molecular phylogeny of Pteris is presented for 135 species, based on cpDNA rbcL and matK and using maximum parsimony, maximum likelihood and Bayesian inference approaches. The inferred phylogeny was used to assess the biogeographical history of Pteris and to reconstruct the evolution of one ecological and four morphological characters commonly used for infrageneric classifications.

Key Results

The monophyly of Pteris remains uncertain, especially regarding the relationship of Pteris with Actiniopteris + Onychium and Platyzoma. Pteris comprises 11 clades supported by combinations of ecological and morphological character states, but none of the characters used in previous classifications were found to be exclusive synapomorphies. The results indicate that Pteris diversified around 47 million years ago, and when species colonized new geographical areas they generated new lineages, which are associated with morphological character transitions.

Conclusions

This first phylogeny of Pteris on a global scale and including more than half of the diversity of the genus should contribute to a new, more reliable infrageneric classification of Pteris, based not only on a few morphological characters but also on ecological traits and geographical distribution. The inferred biogeographical history highlights long-distance dispersal as a major process shaping the worldwide distribution of the species. Colonization of different niches was followed by subsequent morphological diversification. Dispersal events followed by allopatric and parapatric speciation contribute to the species diversity of Pteris.     

Taxonomy, phylogeny, biogeography, and ecology of Sabulodinium undulatum (Dinophyceae), including an emended description of the species

Samples of Sabulodinium undulatum Saunders et Dodge, the type species of the monospecific genus, were collected and characterized from Germany, Russia, Japan and Canada. This species has a laterally flattened, oval cell with a truncated apex and a dorsally pointing small episome. The dorsal margin of the hyposome has an undulating shape. A dorsal spine and/or antapical spine are sometimes present. The specimens of this heterotrophic species are 27.5–42.5 μm long and 18.5–36.0 μm wide, and have a theca with the plate arrangement apical pore complex (APC) 5′ 1a 6″ 5c 4s 6′′′ 1′′′′. The shape of the dorsal theca is variable. The species distribution seems to be restricted to northern temperate regions. Sabulodinium undulatum occurred in all sandy eulittoral areas throughout the year and was also present in sandy sublittoral and supralittoral (beach) samples. The species can tolerate a broad range of temperatures and salinities. Sabulodinium occurred from −2.0 to +24.3°C and 4–35 salinity. It is likely that the observed variability in morphology and habitat reflect several varieties. We propose to establish three varieties within the species, namely S. undulatum var. undulatum, S. undulatum var. glabromarginatum, and S. undulatum var. monospinum. The systematic position of Sabulodinium is discussed in the context of comparative morphological and molecular phylogenetic data.     

The biogeography of Lessoniaceae,with special reference to Macrocystis C. Agardh (Phaeophyta: Laminariales)

The bipolar distributions of Lessoniaceae and Macrocystis have been explained by migration out of a ‘centre of origin’ and across the tropics by means of dispersal, but controversy centres on the issue: which sector is the true centre of origin? We provide biological, palaeo-oceanographic and geological evidence that leads us to reject the centre of origin/migration theory and methodology in explaining ‘bipolarity’. As an alternative, a process of vicariant differentiation out of a Pacific Ocean/Southern Ocean ancestral complex is proposed. Moreover the biogeographic relationship between the southwest and northeast Pacific is shown to be a general phenomenon and not restricted to marine algae.     

Changing perspectives on the biogeography of the tropical South Pacific: influences of dispersal, vicariance and extinction

Aim The biogeographical patterns and drivers of diversity on oceanic islands in the tropical South Pacific (TSP) are synthesized. We use published studies to determine present patterns of diversity on TSP islands, the likely sources of the biota on these islands and how the islands were colonized. We also investigate the effect of extinctions. Location We focus on oceanic islands in the TSP. Methods We review available literature and published molecular studies. Results Examples of typical island features (e.g. gigantism, flightlessness, gender dimorphism) are common, as are adaptive radiations. Diversity decreases with increasing isolation from mainland sources and with decreasing size and age of archipelagos, corresponding well with island biogeographical expectations. Molecular studies support New Guinea/Malesia, New Caledonia and Australia as major source areas for the Pacific biota. Numerous studies support dispersal‐based scenarios, either over several 100 km (long‐distance dispersal) or over shorter distances by island‐hopping (stepping stones) and transport by human means (hitch‐hiking). Only one vicariance explanation, the eastward drift of continental fragments (shuttles) that may have contributed biota to Fiji from New Caledonia, is supported by some geological evidence, although there is no evidence for the transport of taxa on shuttle fragments. Another vicariance explanation, the existence of a major continental landmass in the Pacific within the last 100 Myr (Atlantis theory), receives little support and appears unlikely. Extinction of lineages in source areas and persistence in the TSP has probably occurred many times and has resulted in misinterpretation of biogeographical data. Main conclusions Malesia has long been considered the major source region for the biota of oceanic islands in the TSP because of shared taxa and high species diversity. However, recent molecular studies have produced compelling support for New Caledonia and Australia as alternative important source areas. They also show dispersal events, and not vicariance, to have been the major contributors to the current biota of the TSP. Past extinction events can obscure interpretations of diversity patterns.     

The phylogeny and biogeography of Gentiana L. sect. Ciminalis (Adans.) Dumort.: A historical interpretation of distribution ranges in the European high mountains

Gentiana sect. Ciminalis consists of seven mostly ecologically or geographically vicariant and closely related species which are distributed throughout the South and Central European high mountains. The analysis of a RAPD data set and trn L-intron and ITS sequences resulted in slightly different phylogenetic hypotheses. In the preferred hypothesis the group consists of two completely resolved main lineages: 1) G. clusii and G. alpina. 2) G. dinarica, G. acaulis, G. ligustica, G. angustifolia and G. occidentalis. The most important conclusions we have drawn from this phylogenetic hypothesis and from the observed patterns of molecular variation are: 1) The calcifuge ecology of G. acaulis and G. alpina evolved independently from calcicole ancestors. 2) Among the calcicole taxa speciation proceeded from East to West in a simple vicariant pattern. 3) The application of a provisional molecular clock indicates that speciation events in sect. Ciminalis probably occurred in the Quaternary. 4) Differences in infraspecific genetic variation among the widespread species suggest that G. alpina probably experienced more recent dispersal or gene flow than G. clusii and G. acaulis. 5) The large number of mutations in the lineage leading to G. angustifolia, compared to the few mutations in the lineage leading to G. dinarica, may be a result of their different population histories. While the extant range of G. angustifolia was strongly affected by Quaternary climatic fluctuations, that of G. dinarica has had a more stable climatic history. 6) The low number of mutations and the basal position in one clade of the preferred cladogram leads to the conclusion that G. dinarica is the species most similar to the last common ancestor of sect. Ciminalis.