Inferring the biogeographic origins of inter‐continental disjunct endemics using a Bayes‐DIVA approach

The arcto‐Tertiary relictual flora is comprised of many genera that occur non‐contiguously in the temperate zones of eastern Asia, Europe, eastern North America, and western North America. Within each distributional area, species are typically endemic and may thus be widely separated from closely related species within the other areas. It is widely accepted that this common pattern of distribution resulted from of the fragmentation of a once more‐continuous arcto‐Tertiary forest. The historical biogeographic events leading to the present‐day disjunction have often been investigated using a phylogenetic approach. Limitations to these previous studies have included phylogenetic uncertainty and uncertainty in ancestral range reconstructions. However, the recently described Bayes‐DIVA method handles both types of uncertainty. Thus, we used Bayes‐DIVA analysis to reconstruct the stem lineage distributions for 185 endemic lineages from 23 disjunct genera representing 17 vascular plant families. In particular, we asked whether endemic lineages within each of the four distributional areas more often evolved from (1) widespread ancestors, (2) ancestors dispersed from other areas, or (3) endemic ancestors. We also considered which of these three biogeographic mechanisms may best explain the origins of arcto‐Tertiary disjunct endemics in the neotropics. Our results show that eastern Asian endemics more often evolved from endemic ancestors compared to endemics in Europe and eastern and western North America. Present‐day endemic lineages in the latter areas more often arose from widespread ancestors. Our results also provide anecdotal evidence for the importance of dispersal in the biogeographic origins of arcto‐Tertiary species endemic in the neotropics.     

Phylogeographic history of the woodwardioid ferns,including species from the Himalayas

The woodwardioid ferns are well-represented in the Northern Hemisphere, where they are disjunctly distributed throughout the warm temperate and subtropical regions of North America, Europe, and Asia. To infer the biogeographic history of the woodwardioid ferns, the phylogeny of Woodwardia was estimated using rbcL and rps4 sequences from divergent distribution regions including the Himalayas. Phylogenetic results support Woodwardia as a monophyletic group with Woodwardia areolatae and W. virginica as basal, these two species from eastern North America diverged early, which are sister clades to the remaining species from America, Europe, and Asia. Based on analyses of the fossil records of these species for divergence times, Woodwardia species were estimated to have diverged initially in the Paleogene of North America. After its New World origin, a greater diversification and expansion of Woodwardia occurred in eastern Eurasia, with the European arrival of Woodwardia radicans during the Middle Miocene. Compared to earlier reports, a migration back into North America via the Bering land bridge is consistent with these data.     

Construction of the phylogenetic model for the genera of the tribe Arctiini (Lepidoptera,Arctiidae) with the SYNAP method

A new scheme of the phylogeny of the tribe Arctiini is proposed. The Western Mediterranean genus Atlantarctia is considered the most primitive one in the tribe; the rest of genera form two large clades Arctia-Pericallia and Gonerda-Platyprepia. The first clade is supposed to have been subjected to radiation in western Eurasia, and the second clade, in Asia and North America in the Palaeogene when the eastern part of Asia was isolated from western Eurasia. Subsequently, most probably in the Neogene-Pleistocene, representatives of both clades spread over the whole Eurasia and North America. The Arctiini fauna of the tundra zone, which includes the genera Acerbia and Pararctia, was formed in Asia and North America, whereas the subboreal fauna (both steppe and nemoral) originated in western Eurasia. The boreal genus Borearctia has most likely also originated in Asia.     

Placing the origin of two species-rich genera in the late cretaceous with later species divergence in the tertiary: a phylogenetic, biogeographic and molecular dating analysis of Piper and Peperomia (Piperaceae)

Nearly all of the species diversity in Piperaceae is encompassed within Piper and Peperomia. Both genera are pan-tropical with areas of diversification in the Neotropics and Southeast Asia. Piperaceae are less diverse in Africa with only two native species of Piper. This study examines the distribution of both Piper and Peperomia with representative samples from the Neotropics, Asia, Pacific Islands, and Africa. Molecular dating is used to place an age for the crown clades of Piper and Peperomia as well as ages for diversification within the clades. Both genera have origins in the late Cretaceous, but species level diversification occurred much later in the Tertiary. Biogeography of both genera are correlated with paleoclimate evidence to better explain the distribution and diversification of these large genera.     

Multigene Molecular Phylogeny and Biogeographic Diversification of the Earth Tongue Fungi in the Genera Cudonia and Spathularia (Rhytismatales,Ascomycota)

The family Cudoniaceae (Rhytismatales, Ascomycota) was erected to accommodate the “earth tongue fungi” in the genera Cudonia and Spathularia. There have been no recent taxonomic studies of these genera, and the evolutionary relationships within and among these fungi are largely unknown. Here we explore the molecular phylogenetic relationships within Cudonia and Spathularia using maximum likelihood and Bayesian inference analyses based on 111 collections from across the Northern Hemisphere. Phylogenies based on the combined data from ITS, nrLSU, rpb2 and tef-1α sequences support the monophyly of three main clades, the /flavida, /velutipes, and /cudonia clades. The genus Cudonia and the family Cudoniaceae are supported as monophyletic groups, while the genus Spathularia is not monophyletic. Although Cudoniaceae is monophyletic, our analyses agree with previous studies that this family is nested within the Rhytismataceae. Our phylogenetic analyses circumscribes 32 species-level clades, including the putative recognition of 23 undescribed phylogenetic species. Our molecular phylogeny also revealed an unexpectedly high species diversity of Cudonia and Spathularia in eastern Asia, with 16 (out of 21) species-level clades of Cudonia and 8 (out of 11) species-level clades of Spathularia. We estimate that the divergence time of the Cudoniaceae was in the Paleogene approximately 28 Million years ago (Mya) and that the ancestral area for this group of fungi was in Eastern Asia based on the current data. We hypothesize that the large-scale geological and climatic events in Oligocene (e.g. the global cooling and the uplift of the Tibetan plateau) may have triggered evolutionary radiations in this group of fungi in East Asia. This work provides a foundation for future studies on the phylogeny, diversity, and evolution of Cudonia and Spathularia and highlights the need for more molecular studies on collections from Europe and North America.     

Phylogeny and Biogeography of Thuja L. （Cupressaceae）, an Eastern Asian and North American Disjunct Genus

In order to develop better insights into biogeographic patterns of eastern Asian and North American disjunct plant genera, sequences of nuclear ribosomal DNA internal transcribed spacer (nr DNAITS) region were used to estimate interspecific relationships of Thuja L. (Cupressaceae) and infer its biogeography based on the phylogeny. According to the phylogenetic analysis, two clades were recognized. The first clade included Thuja plicata D. Don (western North America) and T. koraiensis Nakai (northeastern Asia), and the second one contained T. occidentalis (Gord.) Cart. (Japan). The ancestral area of Thuja was inferred to be eastern Asia, and two dispersal events were responsible for the modern distribution of Thuja in North America. Both the North Atlantic land bridge and Bering land bridge were possible routes for the migration of ancestral populations to North America.     

Molecular phylogeny of the harvestmen genus Sabacon (Arachnida: Opiliones: Dyspnoi) reveals multiple Eocene–Oligocene intercontinental dispersal events in the Holarctic

We investigated the phylogeny and biogeographic history of the Holarctic harvestmen genus Sabacon, which shows an intercontinental disjunct distribution and is presumed to be a relatively old taxon. Molecular phylogenetic relationships of Sabacon were estimated using multiple gene regions and Bayesian inference for a comprehensive Sabacon sample. Molecular clock analyses, using relaxed clock models implemented in BEAST, are applied to date divergence events. Biogeographic scenarios utilizing S-DIVA and Lagrange C++ are reconstructed over sets of Bayesian trees, allowing for the incorporation of phylogenetic uncertainty and quantification of alternative reconstructions over time. Four primary well-supported subclades are recovered within Sabacon: (1) restricted to western North America; (2) eastern North American S. mitchelli and sampled Japanese taxa; (3) a second western North American group and taxa from Nepal and China; and (4) eastern North American S. cavicolens with sampled European Sabacon species. Three of four regional faunas (wNA, eNA, East Asia) are thereby non-monophyletic, and three clades include intercontinental disjuncts. Molecular clock analyses and biogeographic reconstructions support nearly simultaneous intercontinental dispersal coincident with the Eocene–Oligocene transition. We hypothesize that biogeographic exchange in the mid-Tertiary is likely correlated with the onset of global cooling, allowing cryophilic Sabacon taxa to disperse within and among continents. Morphological variation supports the divergent genetic clades observed in Sabacon, and suggests that a taxonomic revision (e.g., splitting Sabacon into multiple genera) may be warranted.     

Species richness in plant genera disjunct between temperate eastern Asia and North America

The species richness of 109 amphi-Pacific disjunct genera was examined in eastern Asia and North America. Although the entire flora of eastern Asia contains approximately one-third more species than that of North America, the difference in species richness among disjunct taxa is less. When woody and herbaceous genera are considered separately, the former exhibit a strong diversity bias favouring eastern Asia whereas there is no significant difference in diversity between continents among herbaceous genera. This result is not due to habitat differences between woody and herbaceous genera, because the disjunct herbs inhabit primarily moist forests and woodlands. This result is also not related to relative phylogenetic advancement, even though older major lineages of plants tend to have a predominance of woody taxa. Woody genera are distributed in lower latitudes than herbaceous genera on both continents, and both woody and herbaceous genera are distributed in lower latitudes in eastern Asia than in North America. The North American temperate flora is primarily a relict of a flora form 7 more widespread throughout the Northern Hemisphere. Contemporary patterns of diversity suggest that the effects of climate changes in the late Tertiary were less severe in eastern Asia and promoted diversification, but were more severe in North America and may have caused widespread extinction. The difference in the effect of climate change on diversity in herbaceous and woody lineages reflects the different ecological relationships of species having these contrasting life forms. Clearly, the contemporary floras of eastern Asia and North America bear the imprint of history and emphasize the important interface between ecological relationships and evolutionary responses.     

Phylogeny and biogeography of Rhododendron subsection Pontica, a group with a tertiary relict distribution

Rhododendron subgenus Hymenanthes subsection Pontica is exceptional among Tertiary relict groups in having a high proportion of species (4 of 11) native to SW Eurasia. A phylogeny based on cpDNA matK and trnL-F indicated that multiple Pontica lineages colonised each of SW Eurasia, SE North America, and NE Asia, with little or no speciation within regions thereafter. Therefore, multiple (3-4) Pontica lineages survived the Quaternary in SW Eurasia, in contrast to other Tertiary relict genera. Pontica comprises two major clades, one of which is wholly Eurasian, and paraphyletic with respect to at least some of the remaining 200 species of subgenus Hymenanthes, which are all distributed in SE Asia. The other clade has species from W and SE North America, SW Eurasia, and NE Asia. According to synonymous matK substitution data, the two clades diverged 9-6 million years ago (mya), whereas most divergence within them happened 5-3 mya. Although the phylogeny indicates probable trans-Atlantic migration for one of two America-Eurasia disjunctions in Pontica, the timing supports migration via Beringia for both.     

Phylogenetic relationships among New World Scrophularia L. (Scrophulariaceae): new insights inferred from DNA sequence data

The genus Scrophularia L. (Scrophulariaceae) comprises 200?C300 species, of which approximately 19 are distributed in North America and the Greater Antilles. To investigate phylogenetic and biogeographic relationships of the New World species, two intergenic spacers (trnQ-rps16 and psbA-trnH) of chloroplast DNA and nuclear ribosomal ITS were sequenced. Phylogenetic analyses revealed three distinct New World clades that correspond to their geographical distribution and are corroborated by morphological characters. Phylogenetic inference indicates the eastern American S. marilandica L. as sister to all Antillean species; for colonization of the Caribbean archipelago, a late Miocene dispersal event from the North American mainland is assumed. There is evidence for a hybrid origin of the most widespread North American species, S. lanceolata Pursh. The results further suggest that S. nodosa L. is sister to all New World and three Japanese species of Scrophularia. The latter form an Eastern Asian?CEastern North American (EA-ENA) disjunction with six New World species. We propose an eastern Asian origin for the New World taxa of Scrophularia. Divergence times estimated using a relaxed molecular clock model suggest one or more Miocene migration events from eastern Asia to the New World via the Bering Land Bridge followed by diversification in North America.     

Historical biogeography and phenotype‐phylogeny of Chroodiscus (lichenized Ascomycota: Ostropales: Graphidaceae)

Aim To analyse the historical biogeography of the lichen genus Chroodiscus using a phenotype‐based phylogeny in the context of continental drift and evolution of tropical rain forest vegetation. Location All tropical regions (Central and South America, Africa, India, Southeast Asia, north‐east Australia). Methods We performed a phenotype‐based phylogenetic analysis and ancestral character state reconstruction of 14 species of the lichen genus Chroodiscus, using paup * and mesquite ; dispersal–vicariance analysis (DIVA) and dispersal–extinction–cladogenesis (DEC) modelling to trace the geographical origin of individual clades; and ordination and clustering by means of pc‐ord , based on a novel similarity index, to visualize the biogeographical relationships of floristic regions in which Chroodiscus occurs. Results The 14 species of Chroodiscus show distinctive distribution patterns, with one pantropical and one amphi‐Pacific taxon and 12 species each restricted to a single continent. The genus comprises four clades. DIVA and DEC modelling suggest a South American origin of Chroodiscus in the mid to late Cretaceous (120–100 Ma), with subsequent expansion through a South American–African–Indian–Southeast Asian–Australian dispersal route and late diversification of the argillaceus clade in Southeast Asia. Based on the abundance of extant taxa, the probability of speciation events in Chroodiscus is shown to be extremely low. Slow dispersal of foliicolous rain forest understorey lichens is consistent with estimated phylogenetic ages of individual species and with average lengths of biological species intervals in fungi (10–20 Myr). Main conclusions The present‐day distribution of Chroodiscus can be explained by vicariance and mid‐distance dispersal through the interconnection or proximity of continental shelves, without the need for recent, trans‐oceanic long‐distance dispersal. Phylogenetic reconstruction and age estimation for Chroodiscus are consistent with the ‘biotic ferry’ hypothesis: a South American origin and subsequent eastward expansion through Africa towards Southeast Asia and north‐eastern Australia via the Indian subcontinent. The present‐day pantropical distributions of many clades and species of foliicolous lichens might thus be explained by eastward expansion through continental drift, along with the evolution of modern rain forests starting 120 Ma, rather than by the existence of a hypothetical continuous area of pre‐modern rain forest spanning South America, Africa and Southeast Asia during the mid and late Cretaceous.     

Phylogeny, biogeography, and molecular dating of cornelian cherries (Cornus, Cornaceae): tracking Tertiary plant migration

Data from four DNA regions (rbcL, matK, 26S rDNA, and ITS) as well as extant and fossil morphology were used to reconstruct the phylogeny and biogeographic history of an intercontinentally disjunct plant group, the cornelian cherries of Cornus (dogwoods). The study tests previous hypotheses on the relative roles of two Tertiary land bridges, the North Atlantic land bridge (NALB) and the Bering land bridge (BLB), in plant migration across continents. Three approaches, the Bayesian, nonparametric rate smoothing (NPRS), and penalized likelihood (PL) methods, were employed to estimate the times of geographic isolations of species. Dispersal and vicariance analysis (DIVA) was performed to infer the sequence and directionality of biogeographic pathways. Results of phylogenetic analyses suggest that among the six living species, C. sessilis from western North America represents the oldest lineage, followed by C. volkensii from Africa. The four Eurasian species form a clade consisting of two sister pairs, C. mas-C. officinalis and C. chinensis-C. eydeana. Results of DIVA and data from fossils and molecular dating indicate that the cornelian cherry subgroup arose in Europe as early as the Paleocene. Fossils confirm that the group was present in North America by the late Paleocene, consistent with the DIVA predictions that, by the end of the Eocene, it had diversified into several species and expanded its distribution to North America via the NALB and to Africa via the last direct connection between Eurasia and Africa prior to the Miocene, or via long-distance dispersal. The cornelian cherries in eastern Asia appear to be derived from two independent dispersal events from Europe. These events are inferred to have occurred during the Oligocene and Miocene. This study supports the hypothesis that the NALB served as an important land bridge connecting the North American and European floras, as well as connecting American and African floras via Europe during the early Tertiary.     

Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera,Nymphalidae, Satyrinae)

Peña, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. (2010). Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae).—Zoologica Scripta, 39, 243–258. The diverse butterfly subtribe Euptychiina was thought to be restricted to the Americas. However, there is mounting evidence for the Oriental Palaeonympha opalina being part of Euptychiina and thus a disjunct distribution between it (in eastern Asia) and its sister taxon (in eastern North America). Such a disjunct distribution in both eastern Asia and eastern North America has never been reported for any butterfly taxon. We used 4447 bp of DNA sequences from one mitochondrial gene and four nuclear genes for 102 Euptychiina taxa to obtain a phylogenetic hypothesis of the subtribe, estimate dates of origin and diversification for major clades and perform a biogeographic analysis. Euptychiina originated 31 Ma in South America. Early Euptychiina dispersed from North to South America via the temporary connection known as GAARlandia during Eocene–Oligocene times. The current disjunct distribution of the Oriental Palaeonympha opalina is the result of a northbound dispersal of a lineage from South America into eastern Asia via North America. The common ancestor of Palaeonympha and its sister taxon Megisto inhabited the continuous forest belt across North Asia and North America, which was connected by Beringia. The closure of this connection caused the split between Palaeonympha and Megisto around 13 Ma and the severe extinctions in western North America because of the climatic changes of the Late Miocene (from 13.5 Ma onwards) resulted in the classic ‘eastern Asia and eastern North America’ disjunct distribution.     

Phylogeny and Biogeography of Thuja L. (Cupressaceae), an Eastern Asian and North American Disjunct Genus

In order to develop better insights into biogeographic patterns of eastern Asian and North American disjunct plant genera, sequences of nuclear ribosomal DNA internal transcribed spacer (nr DNA ITS) region were used to estimate interspecific relationships of Thuja L. (Cupressaceae) and infer its biogeography based on the phylogeny. According to the phylogenetic analysis, two clades were recognized. The first clade included Thuja plicata D. Don (western North America) and T. koraiensis Nakai (northeastern Asia), and the second one contained T. occidentalis (Gord.) Carr. (Japan). The ancestral area of Thuja was inferred to be eastern Asia, and two dispersal events were responsible for the modern distribution of Thuja in North America. Both the North Atlantic land bridge and Bering land bridge were possible routes for the migration of ancestral populations to North America.     

Molecular phylogeny of Anaphalis (Asteraceae, Gnaphalieae) with biogeographic implications in the Northern Hemisphere

Anaphalis is the largest Asian genus in the tribe Gnaphalieae (Asteraceae) and has its greatest species diversity in the eastern Himalayas. The nuclear internal and external transcribed spacers were sequenced for Anaphalis species, with an emphasis on the eastern Himalayan taxa to examine the monophyly and construct the phylogenetic relationships of and within the genus. The results suggest that all species of Anaphalis are nested with Helichrysum, showing a close relationship with a Mediterranean–Asian group of Helichrysum. Although the monophyly of Anaphalis is only weakly supported, two clades within the genus are well recognized, each consisting of two subgroups. The inferred phylogenetic relationships within Anaphalis correspond to the shape of leaf base, rather than the morphology of the capitula and phyllaries that are usually used for species delimitation and classification in the genus. All four subgroups of Anaphalis are common and diversified in the eastern Himalayas with multiple dispersals out of this region. The sole North American species of Anaphalis is best hypothesized to be the result of long-distance dispersal or overland migration via Bering land bridge from Asia. Our analyses suggest that the extant distribution of Anaphalis has most likely resulted one radiation into the eastern Himalayas followed by repeated independent dispersals and/or radiations mostly into eastern Asia but also into the western Himalayas, North America, and southeast Asia.     

Evolutionary history of Gymnocarpos (Caryophyllaceae) in the arid regions from North Africa to Central Asia

Gymnocarpos has only about ten species distributed in the arid regions of Asia and Africa, but it exhibits a geographical disjunction between eastern Central Asia and western North Africa and Minor Asia. We sampled eight species of the genus and sequenced two chloroplast regions (rps16 and psbB–psbH), and the nuclear rDNA (ITS) to study the phylogeny and biogeography. The results of the phylogenetic analyses corroborated that Gymnocarpos is monophyletic, in the phylogenetic tree two well supported clades are recognized: clade 1 includes Gymnocarpos sclerocephalus and G. decandrus, mainly the North African group, whereas clade 2 comprises the remaining species, mainly in the Southern Arabian Peninsula. Molecular dating analysis revealed that the divergence age of Gymnocarpos was c. 31.33 Mya near the Eocene and Oligocene transition boundary, the initial diversification within Gymnocarpos dated to c. 6.69 Mya in the late Miocene, and the intraspecific diversification mostly occurred during the Quaternary climate oscillations. Ancestral area reconstruction suggested that the Southern Arabian Peninsula was the ancestral area for Gymnocarpos. Our conclusions revealed that the aridification since mid‐late Miocene significantly affected the diversification of the genus in these areas.     

Phylogenetic relationships in Gleditsia (Leguminosae) based on ITS sequences

We used nucleotide sequences from the internal transcribed spacers and 5.8S gene of nuclear ribosomal DNA to test competing phylogenetic and biogeographic hypotheses in Gleditsia. Eleven of 13 Gleditsia species were sampled, along with two species of its sister genus, Gymnocladus. Analyses of ITS data and of a combined data set that included sequences of ITS and two chloroplast genes supported several conclusions that were interpreted in light of fossil data and current legume phylogeny. Gleditsia and Gymnocladus appear to have originated in eastern Asia during the Eocene. Eastern North American species of both genera most likely evolved from ancestors that migrated across the Bering land bridge, but the eastern Asian/eastern North American disjunction appears to be much older in Gymnocladus than in Gleditsia. Gleditsia amorphoides, from temperate South America, is sister to the rest of the genus, suggesting early long-distance dispersal from Asia. The remainder of Gleditsia is divided into three unresolved clades, possibly indicating a split early in the evolution of the genus. Two of those clades contain only Asian species, and one contains Asian and North American species. The North American species, Gleditsia triacanthos and Gleditsia aquatica, are polymorphic and paraphyletic with respect to their ITS and cpDNA sequences, which suggests recent diversification.     

Dispersal vs. vicariance in the Mediterranean: historical biogeography of the Palearctic Pachydeminae (Coleoptera, Scarabaeoidea)

Aim The geological evolution of the Mediterranean region is largely the result of the Tertiary collision of the African and Eurasian Plates, but also a mosaic of migrating island arcs, fragmenting tectonic belts, and extending back‐arc basins. Such complex paleogeography has resulted in a ‘reticulate’ biogeographical history, in which Mediterranean biotas repeatedly fragmented and merged as dispersal barriers appeared and disappeared through time. In this study, dispersal‐vicariance analysis (DIVA) is used to assess the relative role played by dispersal and vicariance in shaping distribution patterns in the beetle subfamily Pachydeminae Reitter, 1902 (Scarabaeoidea), an example of east–west Mediterranean disjunction. Location The Mediterranean region, including North Africa, the western Mediterranean, Balkans–Anatolia, Middle East, Caucasus, the Iranian Plateau, and Central Asia. Methods A phylogenetic hypothesis of the Palearctic genera of Pachydeminae in conjunction with distributional data was analysed using DIVA. This method reconstructs the ancestral distribution in a given phylogeny based on the vicariance model, while allowing dispersal and extinction to occur. Unlike other methods, DIVA does not enforce area relationships to conform to a hierarchical ‘area cladogram’, so it can be used to reconstruct ‘reticulate’ biogeographical scenarios. Results Optimal reconstructions, requiring 23 dispersal events, suggest that the ancestor of Pachydeminae was originally present in the south‐east Mediterranean region. Basal splitting within the subfamily was caused by vicariance events related to the late Tertiary collision of the African microplates Apulia and Arabia with Eurasia, and the resultant arise of successive dispersal barriers (e.g. the Red Sea, the Zagros Mountains). Subsequent diversification in Pachydeminae involved multiple speciation events within the Middle East and Iran–Afghanistan regions, which gave rise to the least speciose genera of Pachydeminae (e.g. Otoclinius Brenske, 1896). Finally, the presence of Pachydeminae in the western Mediterranean region seems to be the result of a recent dispersal event. The ancestor of the Iberian genera Ceramida Baraud, 1987 and Elaphocera Gené, 1836 probably dispersed from the Middle East to the Iberian Peninsula across North Africa and the Gibraltar Strait during the ‘Messinian salinity crisis’ at the end of the Miocene. Main conclusions Although the basal diversification of Pachydeminae around the Mediterranean appears to be related to vicariance events linked to the geological formation of the Mediterranean Basin, dispersal has also played a very important role. Nearly 38% of the speciation events in the phylogeny resulted from dispersal to a new area followed by allopatric speciation between lineages. Relationships between western and eastern Mediterranean disjuncts are usually explained by dispersal through Central Europe. The biogeographical history of the Pachydeminae corroborates other biogeographical studies that consider North Africa to be an alternative dispersal route by which Mediterranean taxa could have achieved circum‐Mediterranean distributions.     

几种石耳科地衣生物地理学关系的分子评估(英文)

对具有北美东部和亚洲东部近缘生物地理学关系的大型地衣种类进行系统发育检测较少。笔者获得了石耳科中8个种10个样本nrDNA的ITS完整序列,并对本科中在分布上具有重要意义的4个种的生物地理学关系进行了分析和评估。通过对细胞核核糖体DNA变异性较强的ITS段碱基序列的比较分析,支持放射盘石耳(Umbilicaria muehlenbergii)、宾州疱脐衣(Lasallia pennsylvanica)为同种型间断分布,美味石耳(U.escu-lenta)和大叶石耳(U.mammulata)间已经达到了相当高的分化程度,为异种型间断分布。地理学分析显示该两种石耳的分化可能是由于太平洋的扩展和隔离所造成的结果。同种型间断分布种类的不同地区的样本之间的差异较小。推测了不同类型间断分布种类的分化时间,美味石耳与大叶石耳的分化发生在中新世(23～5.33 Mya)中期,白令海峡形成期间。     

Plastomic data shed new light on the phylogeny,biogeography, and character evolution of the family Crassulaceae

Crassulaceae is a mid-sized family of angiosperms, most species of which are herbaceous succulents, usually with 5-merous flowers and one or two whorls of stamens. Although previous phylogenetic studies revealed seven major “clades” in Crassulaceae and greatly improved our understanding of the evolutionary history of the family, relationships among major clades are still contentious. In addition, the biogeographic origin and evolution of important morphological characters delimiting infrafamilial taxa have not been subject to formal biogeographic and character evolution analyses based on a well-supported phylogeny backbone. In this study, we used plastomic data of 52 species, representing all major clades revealed in previous studies to reconstruct a robust phylogeny of Crassulaceae, based on which we unraveled the spatiotemporal framework of diversification of the family. We found that the family may originate in southern Africa and then dispersed to the Mediterranean, from there to eastern Asia, Macaronesia, and North America. The crown age of Crassulaceae was dated at ca. 63.93 million years ago, shortly after the Cretaceous–Paleogene (K-Pg) boundary. We also traced the evolution of six important morphological characters previously used to delimit infrafamilial taxa and demonstrated widespread parallel and convergent evolution of both vegetative (life form and phyllotaxis) and floral characters (number of stamen whorls, petals free or fused, and flower merism). Our results provide a robust backbone phylogeny as a foundation for further investigations, and also some important new insights into biogeography and evolution of the family Crassulaceae.