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, ●● , ●●–●●.     

Integrated Fossil and Molecular Data Reveal the Biogeographic Diversification of the Eastern Asian-Eastern North American Disjunct Hickory Genus (Carya Nutt.)

The hickory genus (Carya) contains ca. 17 species distributed in subtropical and tropical regions of eastern Asia and subtropical to temperate regions of eastern North America. Previously, the phylogenetic relationships between eastern Asian and eastern North American species of Carya were not fully confirmed even with an extensive sampling, biogeographic and diversification patterns had thus never been investigated in a phylogenetic context. We sampled 17 species of Carya and 15 species representing all other genera of the Juglandaceae as outgroups, with eight nuclear and plastid loci to reconstruct the phylogeny of Carya. The phylogenetic positions of seven extinct genera of the Juglandaceae were inferred using morphological characters and the molecular phylogeny as a backbone constraint. Divergence times within Carya were estimated with relaxed Bayesian dating. Biogeographic analyses were performed in DIVA and LAGRANGE. Diversification rates were inferred by LASER and APE packages. Our results support two major clades within Carya, corresponding to the lineages of eastern Asia and eastern North America. The split between the two disjunct clades is estimated to be 21.58 (95% HPD 11.07-35.51) Ma. Genus-level DIVA and LAGRANGE analyses incorporating both extant and extinct genera of the Juglandaceae suggested that Carya originated in North America, and migrated to Eurasia during the early Tertiary via the North Atlantic land bridge. Fragmentation of the distribution caused by global cooling in the late Tertiary resulted in the current disjunction. The diversification rate of hickories in eastern North America appeared to be higher than that in eastern Asia, which is ascribed to greater ecological opportunities, key morphological innovations, and polyploidy.     

Phylogenetic analysis of eastern Asian and eastern North American disjunct Lespedeza (Fabaceae) inferred from nuclear ribosomal ITS and plastid region sequences

Lespedeza (tribe Desmodieae, Fabaceae) follows a disjunct distribution in eastern Asia and eastern North America. Phylogenetic relationships among its species and related taxa were inferred from nuclear ribosomal internal transcribed spacer (ITS) and plastid sequences (trnH‐psbA, psbK‐psbI, trnK‐matK and rpoC1). We examined 35 species of Lespedeza, two of Kummerowia and one of Campylotropis, the sole constituents of the Lespedeza group. An analysis of these data revealed that the genus Campylotropis is sister to the other two genera. However, we were unable to resolve the relationships between Kummerowia and Lespedeza in the strict consensus trees of parsimony analyses based on plastid and combined DNA data. In the genus Lespedeza, the Old World subgenus Macrolespedeza is monophyletic, whereas the transcontinental subgenus Lespedeza is paraphyletic. Monophyly of eastern Asian species and of North American species is strongly supported. Although inconsistent with the traditional classification, this phylogenetic finding is consistent with seedling morphology. Three subgroups recognized in subgenus Macrolespedeza were unresolved in our phylogenetic trees. An incongruence length difference (ILD) test indicated that the two partitions (nuclear ITS and plastid sequences) were significantly incongruent, perhaps because of hybridization between species in Lespedeza. Most of the primary clades of tribe Desmodieae are Asian, implying that the relatively few New World ones, such as those in Lespedeza, are more recently derived from Asia. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 221–235.     

Phylogenomics,biogeography, and evolution of morphology and ecological niche of the eastern Asian–eastern North American Nyssa (Nyssaceae)

Nyssa (Nyssaceae, Cornales) represents a classical example of the well‐known eastern Asian–eastern North American floristic disjunction. The genus consists of three species in eastern Asia, four species in eastern North America, and one species in Central America. Species of the genus are ecologically important trees in eastern North American and eastern Asian forests. The distribution of living species and a rich fossil record of the genus make it an excellent model for understanding the origin and evolution of the eastern Asian–eastern North American floristic disjunction. However, despite the small number of species, relationships within the genus have remained unclear and have not been elucidated using a molecular approach. Here, we integrate data from 48 nuclear genes, fossils, morphology, and ecological niche to resolve species relationships, elucidate its biogeographical history, and investigate the evolution of morphology and ecological niches, aiming at a better understanding of the well‐known EA–ENA floristic disjunction. Results showed that the Central American (CAM) Nyssa talamancana was sister to the remaining species, which were divided among three, rapidly diversified subclades. Estimated divergence times and biogeographical history suggested that Nyssa had an ancestral range in Eurasia and western North America in the late Paleocene. The rapid diversification occurred in the early Eocene, followed by multiple dispersals between and within the Erasian and North American continents. The genus experienced two major episodes of extinction in the early Oligocene and end of Neogene, respectively. The Central American N. talamancana represents a relic lineage of the boreotropical flora in the Paleocene/Eocene boundary that once diversified in western North America. The results supported the importance of both the North Atlantic land bridge and the Bering land bridge (BLB) for the Paleogene dispersals of Nyssa and the Neogene dispersals, respectively, as well as the role of Central America as refugia of the Paleogene flora. The total‐evidence‐based dated phylogeny suggested that the pattern of macroevolution of Nyssa coincided with paleoclimatic changes. We found a number of evolutionary changes in morphology (including wood anatomy and leaf traits) and ecological niches (precipitation and temperature) between the EA–ENA disjunct, supporting the ecological selection driving trait evolutions after geographic isolation. We also demonstrated challenges in phylogenomic studies of lineages with rapid diversification histories. The concatenation of gene data can lead to inference of strongly supported relationships incongruent with the species tree. However, conflicts in gene genealogies did not seem to impose a strong effect on divergence time dating in our case. Furthermore, we demonstrated that rapid diversification events may not be recovered in the divergence time dating analysis using BEAST if critical fossil constraints of the relevant nodes are not available. Our study provides an example of complex bidirectional exchanges of plants between Eurasia and North America in the Paleogene, but “out of Asia” migrations in the Neogene, to explain the present disjunct distribution of Nyssa in EA and ENA.     

Phylogenetic assessment and biogeographic analyses of tribe <Emphasis Type="Italic">Peracarpeae</Emphasis> (Campanulaceae)

Peracarpeae is a small tribe consisting of three genera: Homocodon, Heterocodon and Peracarpa, with a disjunct distribution between eastern Asia and western North America. Homocodon is endemic to southwestern China and was previously placed in the western North American genus Heterocodon. Our phylogenetic analysis using four plastid markers (matK, atpB, rbcL and trnL-F) suggests the polyphyly of Peracarpeae. Homocodon is sister to a clade consisting of the eastern Asian Adenophora, Hanabusaya and species of Campanula from the Mediterranean region and North America, rather than forming a clade with Heterocodon. Homocodon and its Eurasia relatives are estimated to have diverged in the early Miocene (16.84 mya, 95% HPD 13.35–21.45 mya). The eastern Asian Peracarpa constitutes a clade with the North American Heterocodon, Githopsis and three species of Campanula, supporting a disjunction between eastern Asia and North America in Campanulaceae. The Asian-North American disjunct lineages diverged in the early Miocene (16.17 mya, 95% HPD 13.12–20.9 mya). The biogeographic analyses suggest that Homocodon may be a relict of an early radiation in eastern Asia, and that Peracarpa and its closest North American relatives most likely originated from a Eurasian ancestor.     

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     

Merging Maddenia with the morphologically diverse Prunus (Rosaceae)

Maddenia (Rosaceae) has been distinguished from Prunus on the basis of its tepaloid perianth and one‐ to two‐carpellate gynoecium. These distinctive morphological traits nonetheless overlap with several Prunus spp. Maddenia has previously been shown to be nested within Prunus, more specifically within a clade containing members of subgenera Laurocerasus and Padus, but its phylogenetic position within that clade has not been defined precisely. This study clarifies the position of Maddenia within Prunus through phylogenetic analyses of nuclear ribosomal internal transcribed spacer (ITS) and plastid ndhF sequences, with an expanded sampling of tropical species of subgenus Laurocerasus and the inclusion of three Maddenia spp. The monophyly of Maddenia is supported by both the ITS and ndhF analyses, but both datasets support the inclusion of Maddenia in Prunus. All trees from the ITS analysis and some trees from the ndhF analysis also support a close alliance of Maddenia with a clade comprising temperate species of subgenera Laurocerasus and Padus. On the basis of these results, all recognized species of Maddenia are herein formally transferred to Prunus, which requires four new combinations and one new name: Prunus fujianensis (Y.T.Chang) J.Wen, comb. nov. ; Prunus himalayana J.Wen, nom. nov. ; Prunus hypoleuca (Koehne) J.Wen, comb. nov. ; Prunus hypoxantha (Koehne) J.Wen, comb. nov. ; and Prunus incisoserrata (T.T.Yü & T.C.Ku) J.Wen, comb. nov. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 236–245.     

A rangewide herbarium‐derived dataset indicates high levels of gene flow in black cherry (Prunus serotina)

Isolation by Distance (IBD) is a genetic pattern in which populations geographically closer to one another are more genetically similar to each other than populations which are farther apart. Black cherry (Prunus serotina Ehrh.) (Rosaceae) is a forest tree species widespread in eastern North America, and found sporadically in the southwestern United States, Mexico, and Guatemala. IBD has been studied in relatively few North American plant taxa, and no study has rigorously sampled across the range of such a widespread species. In this study, IBD and overall genetic structure were assessed in eastern black cherry (P. serotina Ehrh. var. serotina), the widespread variety of eastern North America. Eastern North America. Prunus serotina Ehrh. var. serotina (Rosaceae). Dense sampling across the entire range of eastern black cherry was made possible by genotyping 15 microsatellite loci in 439 herbarium samples from all portions of the range. Mantel tests and STRUCTURE analyses were performed to evaluate the hypothesis of IBD and genetic structure. Mantel tests demonstrated significant but weak IBD, while STRUCTURE analyses revealed no clear geographic pattern of genetic groups. The modest geographic/genetic structure across the eastern black cherry range suggests widespread gene flow in this taxon. This is consistent with P. serotina's status as a disturbance‐associated species. Further studies should similarly evaluate IBD in species characteristic of low‐disturbance forests.     

Genetic diversity and distribution patterns of Ecuadorian capuli (Prunus serotina)

Prunus serotina subsp. capuli (Cav.) is an arboreal species with promising economic prospects in the timber, health-food and neutraceutical markets. Despite its cultural and commercial significance, limited information exists with regards to the degree of genetic variation and ecological history of P. serotina in Ecuador. The main objective of this study was to evaluate the degree of genetic diversity and population structure of Ecuadorian P. serotina, as a preliminary step towards understanding the distribution history of this species in Ecuador and establishing germplasm conservation programs. P. serotina samples (217, representing 8 provinces from the Ecuadorian highlands) were characterized using 8 heterologous SSR markers derived from related Prunus species. Expected heterozygosity across samples (H_e = 0.71) reveals a moderate level of genetic diversity for Ecuadorian P. serotina. Nevertheless, simple allele-count analysis indicates that Ecuadorian capuli has a narrower degree of allelic richness relative to collections from the species’ center of origin in North America. Furthermore, pairwise F statistics (0.0069 ≤ Fst ≤ 0.0319) and Nei genetic distance estimates (0.02 ≤ Ds ≤ 0.09) indicate minimal population differentiation within Ecuadorian capuli. However, Bayesian population structure analysis suggests a subtle genetic contrast between germplasm from the Northern and Southern highlands. Certainly, it is of interest to analyze whether this underlying genetic differentiation between the Northern- and Southern-Highland groups is also manifested in morphological, agronomic or other phenotypic characters that could indicate adaptive differences to divergent agro-ecosystems.     

Reticulate evolution in Thuja inferred from multiple gene sequences: implications for the study of biogeographical disjunction between eastern Asia and North America

The eastern Asia-North America disjunction is one of the most interesting biogeographical patterns, but its formation is still in much debate. Here nucleotide sequences of five cpDNA regions, nrDNA ITS and two low-copy nuclear genes (LEAFY, 4CL) were employed to reconstruct the phylogeny and to explore the historical biogeography of Thuja, a typical eastern Asia-North America disjunct genus. High topological discordance was observed between chloroplast and nuclear gene trees, even between different nuclear gene trees, suggesting that Thuja could have a reticulate evolutionary history due to multiple interspecific hybridization events. The eastern Asian species Thuja koraiensis might have obtained its chloroplast genome from the eastern North American species T. occidentalis by chloroplast capture, while the western North American species T. plicata is very likely to have inherited a recombinant cpDNA. Based on the phylogenetic analysis of multiple genes, DIVA-reconstruction of the distribution history, molecular clock estimation and fossil data, we inferred that Thuja could have originated from the high-latitude areas of North America in the Paleocene or earlier with subsequent expansion into eastern Asia through the Bering Land Bridge. The two eastern Asia species T. standishii and T. sutchuenensis have a sister relationship, and their split could have occurred in the Oligocene or early Miocene. In the present study, the selection of molecular markers in biogeographic studies was also discussed. Since most previous studies on the eastern Asia and North America disjunction are based on uniparentally inherited cpDNA and (or) directly sequenced nrDNA ITS data, the historical reticulate evolution in the studied groups might have been underestimated. Therefore, we suggest that multiple genes from different genomes, especially low-copy nuclear genes, be used in this research area in the future.     

Phylogenetic Relationships of Pogoniinae (Vanilloideae, Orchidaceae): An Herbaceous Example of the Eastern North America-Eastern Asia Phytogeographic Disjunction

rb cL DNA sequences, nuclear ribosomal ITS DNA sequences, morphology, and combined evidence. All these matrices produced patterns that agree on the broader Phylogenetic relationship within the clade. Duckeella is sister to all Pogoniinae, South American species of Cleistes are monophyletic, Pogonia is monophyletic and part of a larger clade of temperate taxa (Isotria, Pogonia, and Cleistes divaricata) from North America and Asia. The structure of the cladograms and the high levels of bootstrap support strongly indicate that the genus Cleistes is paraphyletic. The disjunction between tropical South American and temperate North American taxa as well as the disjunction between Pogonia ophioglossoides in eastern North America with P. minor and P. Japonica in eastern Asia are best explained by speciation following a northward longdistance dispersal and subsequent northwestward migration via Bering land bridges in the Tertiary. This phylogenetic study adds an additional herbaceous example to the growing list of plants that demonstrate this classical biogeographic pattern. Received 5 February 1999/ Accepted in revised form 9 June 1999     

Phylogeny and Phytogeography of Eupatorium (Eupatorieae, Asteraceae): Insights from Sequence Data of the nrDNA ITS Regions and cpDNA RFLP

Eupatorium were examined by sequencing the internal transcribed spacers (ITS) of nuclear ribosomal DNA and restriction site analysis of chloroplast DNA. Molecular data provided strong evidence that (1) this genus originated in North America, (2) the genus diverged into three morphological species groups, Eutrochium, Traganthes and Uncasia in North America, and (3) one of the North American Uncasia lineages migrated into temperate Europe and eastern Asia over the Bering land bridge. The estimated divergence times support a late Miocene to early Pliocene migration from North America to Eurasia via the Bering land bridge. A European species was sister to all of the eastern Asian species examined. The disjunct distribution pattern of the genus Eupatorium is incongruent with the classical Arcto-Tertiary geoflora concept. Received 13 September 1999/ Accepted in revised form 4 January 2000     

The role of disjunction and postglacial population expansion on phylogeographical history and genetic diversity of the circumboreal plant Chamaedaphne calyculata

In the last decade a number of studies has illustrated quite different phylogeographical patterns amongst plants with a northern present‐day geographical distribution, spanning the entire circumboreal region and/or circumarctic region and southern mountains. These works, employing several marker systems, have brought to light the complex evolutionary histories of this group. Here I focus on one circumboreal plant species, Chamaedaphne calyculata (leatherleaf), to unravel its phylogeographical history and patterns of genetic diversity across its geographical range. A survey of 29 populations with combined analyses of chloroplast DNA (cpDNA), internal transcribed spacer (ITS) and AFLP markers revealed structuring into two groups: Eurasian/north‐western North American, and north‐eastern North American. The present geographical distribution of C. calyculata has resulted from colonization from two putative refugial areas: east Beringia and south‐eastern North America. The variation of chloroplast DNA (cpDNA) and ITS sequences strongly indicated that the evolutionary histories of the Eurasian/north‐western North American and the north‐eastern North American populations were independent of each other because of a geographical disjunction in the distribution area and ice‐sheet history between north‐eastern and north‐western North America. Mismatch analysis using ITS confirmed that the present‐day population structure is the result of rapid expansion, probably since the last glacial maximum. The AFLP data revealed low genetic diversity of C. calyculata (P = 19.5%, H = 0.085) over the whole geographical range, and there was no evidence of loss of genetic diversity within populations in the continuous range, either at the margins or in formerly glaciated and nonglaciated regions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 761–775.     

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.     

Phylogenetics and biogeography of eastern Asian–North American disjunct genus Pachysandra (Buxaceae) inferred from nucleotide sequences

Abstract Pachysandra is an eastern Asian–North American disjunct genus with three species, two in eastern Asia (Pachysandra axillaris and Pachysandra terminalis) and one in eastern North America (Pachysandra procumbens). Although morphological and cytological studies suggest a close affinity of P. procumbens with P. axillaris, molecular data from nuclear and chloroplast DNA regions have provided conflicting signals. In this study, we tested previous phylogenetic hypotheses using sequences of nuclear ribosomal DNA internal transcribed spacers and chloroplast ndhF gene from multiple individuals of each of the three species. We also estimated the time of divergence between eastern Asia and eastern North America. Our results support the morphological and cytological conclusion that P. procumbens is more closely related to P. axillaris than to P. terminalis. The estimated time of divergence of P. axillaris and P. procumbens was 14.6±5.5 mya, consistent with estimates from many other eastern Asian–North American disjunct genera. The migration of Pachysandra populations from eastern Asia to North America might have occurred by way of the North Atlantic land bridge.     

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.     

Rediscovery of the lost little dogwood Cornus wardiana (Cornaceae)—Its phylogenetic and morphological distinction and implication in the origin of the Arctic‐Sino‐Himalayan disjunction

The dwarf dogwoods (subgenus Arctocrania) have been widely known to consist of three circumboreal species Cornus suecica, Cornus canadensis, and Cornus unalaschkensis. A fourth putative species was discovered from the northern Myanmar in 1937, but it had never been formally reported on. Here, we formally report the species on the basis of phylogenetic and morphological evidence and name it Cornus wardiana Rushforth & Wahlsteen (sp. nov.). We conducted phylogenetic and morphometric analyses to determine its evolutionary relationship and differentiation from the existing relatives. We dated the phylogeny using molecular data and conducted a biogeographic analysis to gain insights into the evolution and biogeography of the Arctic‐Sino‐Himalayan disjunction. The phylogenetic analysis used sequences of the nrITS and plastid matK and rbcL genes and included all four dwarf dogwoods and 20 other species representing the three other major lineages of Cornus and the outgroup. The morphometric analyses included 60 populations and 102 specimens of dwarf dogwood, representing the entire range of the subgenus. The results showed that C. wardiana diverged first within subgenus Arctocrania in the Miocene, from a wide‐spread ancestor. Results from principal component analysis and discriminant analysis also showed that the Myanmar samples are well separated from the others. Taken together, these results suggest that the dwarf dogwood lineage split from the big‐bracted dogwoods in Asia or Asia‐western North America during the late Paleocene and spread widely to form a Eurasia‐North America distribution; the Arctic‐Sino‐Himalayan disjunction was the result of southward migration in the Miocene followed by extinction in the intervening highland areas.     

Givetian Stringocephalid brachiopods from eastern Yunnan of Southwest China with notes on global distribution of the family Stringocephalidae

Stringocephalid brachiopods are widely known in the Givetian, but little knowledge is obtained concerning their palaeobiogeographical patterns globally, therefore further studies with new updates and details are required. In this paper, we describe two new stringocephalid brachiopod genera: Yangirostra asiatica n. gen. n. sp. (subfamily Stringocephalinae) and Chinellirostra rara n. gen. n. sp. (subfamily Bornhardtininae), and a new species Stringocephalus sinensis n. sp., together with an indeterminate species Parastringocephalus sp., from the Givetian (late Middle Devonian) of eastern Yunnan, Southwest China. Moreover, with information of the stringocephalides from North Africa, Alaska, North America, Europe and Northeast Eurasia, we compile a dataset of family Stringocephalidae containing 32 genera in 7 subfamilies globally. Based on our data, subfamily Stringocephalinae brachiopods show cosmopolitism and considerably wide distribution from Siberia to the northern Gondwana margins (i.e., North Africa and Australia). Nevertheless, the Boreal Realm and Palaeotethyan Realm are depicted in this paper at the subfamily level, which is much different from the previous palaeobiogeographical schemes in the Givetian. Furthermore, palaeobiogeographical links between Siberia, the Urals and western North America (Alaska, Canada, Nevada and Sonora) are confirmed by diversification of the subfamilies Omoloninae and Rensselandiinae. Whereas in Eurasia (i.e., western and eastern Europe, North and South China), many endemic species of the subfamilies Bornhardtininae and Geranocephalinae are present, as well as the Kaplexinae and Leioseptathyridinae.     

Phylogeny and biogeography of the carnivorous plant family Sarraceniaceae

The carnivorous plant family Sarraceniaceae comprises three genera of wetland-inhabiting pitcher plants: Darlingtonia in the northwestern United States, Sarracenia in eastern North America, and Heliamphora in northern South America. Hypotheses concerning the biogeographic history leading to this unusual disjunct distribution are controversial, in part because genus- and species-level phylogenies have not been clearly resolved. Here, we present a robust, species-rich phylogeny of Sarraceniaceae based on seven mitochondrial, nuclear, and plastid loci, which we use to illuminate this family's phylogenetic and biogeographic history. The family and genera are monophyletic: Darlingtonia is sister to a clade consisting of Heliamphora+Sarracenia. Within Sarracenia, two clades were strongly supported: one consisting of S. purpurea, its subspecies, and S. rosea; the other consisting of nine species endemic to the southeastern United States. Divergence time estimates revealed that stem group Sarraceniaceae likely originated in South America 44-53 million years ago (Mya) (highest posterior density [HPD] estimate = 47 Mya). By 25-44 (HPD = 35) Mya, crown-group Sarraceniaceae appears to have been widespread across North and South America, and Darlingtonia (western North America) had diverged from Heliamphora+Sarracenia (eastern North America+South America). This disjunction and apparent range contraction is consistent with late Eocene cooling and aridification, which may have severed the continuity of Sarraceniaceae across much of North America. Sarracenia and Heliamphora subsequently diverged in the late Oligocene, 14-32 (HPD = 23) Mya, perhaps when direct overland continuity between North and South America became reduced. Initial diversification of South American Heliamphora began at least 8 Mya, but diversification of Sarracenia was more recent (2-7, HPD = 4 Mya); the bulk of southeastern United States Sarracenia originated co-incident with Pleistocene glaciation, <3 Mya. Overall, these results suggest climatic change at different temporal and spatial scales in part shaped the distribution and diversity of this carnivorous plant clade.