Asian North American Disjunct Research Articles

Phylogenomics and historical biogeography of Hydrangeeae (Hydrangeaceae) elucidate the effects of geologic and climatic dynamics on diversification.

Demonstrating the process of transregional biogeography and mechanisms underlying evolutionary radiations is crucial to understanding biological evolution. Here, we use Hydrangeeae (Hydrangeaceae), a tribe with a unique disjunct distribution and complex trait variations, using a solid phylogenetic framework, to investigate how geographical and climatic factors interact with functional traits to trigger plant evolutionary radiations. We constructed the first highly supported and dated phylogenetic framework using 79 protein-coding genes obtained from 81 plastomes, representing 63 species and all major clades, and found that most extant species originated from asynchronous diversification of two lineages undergoing repeated expansion and retraction, at middle and high latitudes of the Northern Hemisphere between East Asia and North America, during the Eocene to Pleistocene (driven by geologic and climatic dynamics). In accordance with these drivers, interactions of flora between central-eastern China and Japan occurred frequently after the Late Tertiary. We found that resource limitation and range fragmentation probably accelerated the diversification of Hydrangeeae, which supports the resource-use hypothesis. Our study sheds light on the evolutionary radiation and assembly of flora within East Asia, and the East Asian-North American disjunction, through integration of phylogenomic and biogeographic data with functional trait and ecological data.

New discovery of Mahonia fossils from the Pliocene of Yunnan, China, and its biogeographical significance

ABSTRACT Mahonia Nuttall (Berberidaceae Jussieu) is an East Asian‒North American disjunct evergreen genus. There are many fossil records of Mahonia from the Eocene to Pleistocene in North America and Europe, but only a few reliable fossils have been reported from East Asia. Compressed fossil leaflets, described here as Mahonia mangbangensis sp. nov. and Mahonia cf. oiwakensis, were collected from the Pliocene Mangbang Formation in Tengchong County, Yunnan Province, southwest China. These new fossil specimens show the same leaf architectural characteristics as those belonging to the Group Orientales of Mahonia, and represent the youngest occurrence of the genus in East Asia. This new fossil discovery extends the stratigraphic range of Mahonia in East Asia and provides new insights into the potential differentiation and migration routes of the Group Orientales in China. Mahonia probably differentiated during the Pliocene in western Yunnan, representing part of the diversification history of this genus in East Asia.

Phylotranscriptomics reveals the complex evolutionary and biogeographic history of the genus Tsuga with an East Asian-North American disjunct distribution

The disjunct distribution between East Asia and North America is one of the best established biogeographic patterns. A robust phylogeny is fundamental for understanding the biogeographic histories of taxa with this distribution pattern. Tsuga (hemlock) is a genus of Pinaceae with a typical intercontinental disjunct distribution in East Asia and eastern and western North America, and its phylogeny has not been completely reconstructed in previous studies. In this study, we reconstructed a highly resolved phylogeny of Tsuga using 881 nuclear genes, 60 chloroplast genes and 23 mitochondrial genes and explored its biogeographic and reticulate evolutionary history. The results of phylogenetic analysis, molecular dating and ancestral area reconstruction indicate that Tsuga very likely originated from North America in the late Oligocene and dispersed from America to East Asia via the Bering Land Bridge during the middle Miocene. In particular, we found complex reticulate evolutionary pattern among the East Asian hemlock species. T. sieboldii possibly originated from hybridization with the ancestor of T. chinensis from mainland China and T. forrestii as the paternal donor and the ancestor of T. diversifolia and T. ulleungensis as the maternal donor. T. chinensis (Taiwan) could have originated by hybridization together with T. sieboldii and then evolved independently after dispersal to the Taiwan Island, subsequently experiencing mitochondrial DNA introgression with T. chinensis from mainland China. Moreover, our study found that T. chinensis from western China is more closely related to T. forrestii than to T. chinensis from eastern China. The nonmonophyletic T. chinensis needs taxonomic reconsideration.

RAD sequencing resolves the phylogeny, taxonomy and biogeography of Trichophoreae despite a recent rapid radiation (Cyperaceae)

Trichophoreae is a nearly cosmopolitan Cyperaceae tribe that contains ~17 species displaying striking variation in size, inflorescence complexity, and perianth morphology. Although morphologically distinct, the status of its three genera (Cypringlea, Oreobolopsis and Trichophorum) is controversial because recent phylogenetic studies have suggested they might not be reciprocally monophyletic. However, previous analyses have shown conflicting topologies and consistently poor support due to an initial rapid diversification of the tribe. We analysed restriction-site associated DNA sequencing (RADseq) data from nearly all species of the clade, combined with five Sanger-based markers (matK, ndhF, rps16, ETS-1f, ITS) sampled extensively within species. This approach allowed us to resolve deep and shallow relationships within Trichophoreae for the first time, despite an anomaly zone spanning several successive short branches that produced considerable gene tree incongruence. Analyses reveal a primary phylogenetic split of the tribe into two clades roughly corresponding to an East Asian-North American disjunction that dates back to the mid-Miocene, with both clades comprised of a mixture of reduced unispicate and larger taxa with highly compound inflorescences. Morphological characters traditionally used in the circumscription of Trichophoreae genera are shown to be homoplasious. Several of these characters correlate best with climatic conditions, with the most reduced species occurring in open habitats at high latitudes and altitudes. Close relatives with highly compound inflorescences are found in temperate or subtropical forest understories. Cypringlea and Oreobolopsis are deeply nested within Trichophorum, and we merge all three genera into a more broadly circumscribed Trichophorum. We also show that Scirpus filipes is another previously unrecognized East Asian species of Trichophorum with highly compound inflorescences.

The world fauna of Synchroidae Lacordaire, 1859 (Coleoptera, Tenebrionoidea, Synchroidae)

Synchroidae Lacordaire, 1859 is a taxonomically and biologically poorly known group. In the present paper, diagnostic characters used to separate genera are analysed and the phylogenetic relationships within this family are preliminarily investigated. Results suggest that the characteristic Synchroa pangu Hsiao, Li, Liu & Pang, 2016 can be removed to establish a new genus, Thescelosynchroa gen. nov. The new combination, T. pangu (Hsiao, Li, Liu & Pang) gen. et comb. nov., is proposed. The definitions of Synchroa Newman, 1838 and Synchroina Fairmaire, 1898 are revised. Moreover, morphological analysis and character comparison also suggest that the familial placement of Mallodrya subaenea Horn, 1888 is questionable. Six species are re-examined and re-diagnosed: Synchroa chinensis Nikitsky, 1999, S. elongatula Nikitsky, 1999, S. formosana Hsiao, 2015, S. melanotoides Lewis, 1895, S. punctata Newman, 1838 and Synchroina tenuipennis Fairmaire, 1898. The male of S. chinensis and the female of S. formosana are described for the first time. Synchroa elongatula and Synchroina tenuipennis are newly recorded from Laos and Indonesia, respectively. We also hypothesize that the Eastern Asian-North American disjunction of Synchroa could be connected to a Mid-Late Tertiary migration of plants via the Bering Land Bridge.

Molecular systematics and biogeography ofWisteriainferred from nucleotide sequences of nuclear and plastid genes

AbstractPrevious molecular phylogenetic studies of Fabaceae indicated that species ofWisteria, an intercontinental disjunct genus between eastern Asia and eastern North America, formed a clade derived from withinCallerya. However, interspecific relationships were not well resolved or supported. In this study, we used sequences of the nuclear ribosomal DNA internal transcribed spacer region and the chloroplast genematKto examine interspecific relationships and explore implications of the phylogeny for the systematics and biogeography ofWisteria. Our results showed thatWisteriawith deciduous leaves and racemose inflorescences formed a strongly supported clade derived from within the paraphyleticCallerya.Afgekiawas also found to be included withinCallerya. Therefore, our data support the merger ofAfgekia,Callerya, andWisteria. The phylogenetic pattern suggested that the deciduousness inWisteriamay be a derived trait likely in response to temperate climate, and the racemose inflorescences in theAfgekia–Callerya–Wisteriaclade may have evolved from panicles. Our study also provided strong support for the sister relationship of the North American and eastern Asian species ofWisteria. In the Asian clade,Wisteria brachybotrysSiebold & Zucc. of Japan was sister to the clade containingW.floribunda(Willd.) DC of Japan and Korea, andW.sinensis(Sims) Sweet of China. However, our data offered weak support for the sister relationship ofW. floribundaandW. sinensis. Our divergence time and biogeographic analyses suggested that the eastern Asian–North American disjunction inWisteriamay have occurred through a dispersal event in the middle Miocene (13.4 Mya) from the Old World to the New World across the Bering land bridge followed by vicariance in the late Miocene (6.8 Mya). This study added another example to the “out of Asia” migration for the eastern Asian–eastern North American disjunction.

Historical biogeography of Eastern Asian–Eastern North American disjunct Melaphidina aphids (Hemiptera: Aphididae: Eriosomatinae) on Rhus hosts (Anacardiaceae)

Intercontinental biotic disjunctions have been documented and analyzed in numerous Holarctic taxa. Patterns previously synthesized for animals compared to plants suggest that the timing of animal disjunctions are mostly Early Tertiary and were generated by migration and vicariance events occurring in the North Atlantic, while plant disjunctions are mostly Mid-Late Tertiary and imply migration and vicariance over Beringia. Melaphidina aphids (Hemiptera: Aphididae: Fordini) exhibit host-alternating life cycles comprising an obligate seasonal shift between Rhus subgenus Rhus species (Anacardiaceae) and mosses (Bryophyta). Similar to their Rhus hosts, melaphidines are distributed disjunctly between Eastern Asia and Eastern North America. We examined evolutionary relationships within Melaphidina to determine the position of the North American lineage, date its divergence from Asian relatives, and compare these results to a previous historical biogeographic study of Rhus. We sampled nine species and three subspecies representing all six genera of Melaphidina. Data included sequences of mitochondrial cytochrome c oxidase subunits I and II+leucine tRNA, cytochrome b, and nuclear elongation factor 1α genes. Phylogenetic analyses (Bayesian, maximum-likelihood, parsimony) of the combined data (3282bp) supported the monophyly of all genera except Nurudea and Schlechtendalia, due to the position of N. ibofushi. While the exact position of the North American Melaphis was not well resolved, there was high support for a derived position within Asian taxa. The divergence of Melaphis from Asian relatives centered on the Eocene–Oligocene boundary (∼33–35Ma), which coincides with closure of Beringian Land Bridge I. This also corresponded to the Asian–North American disjunction previously estimated for subgenus Rhus spp. We suggest the late-Eocene Bering Land Bridge as the most likely migration route for Melaphis ancestors, as was also hypothesized for North American Rhus ancestors. Results for the Melaphidina disjunction depart from the modal pattern in animal lineages, and present a case where insect and host-plant taxa apparently responded similarly to Tertiary climate change.

Phylogenetic assessment and biogeographic analyses of tribe Peracarpeae (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.

Biogeographic disjunction between eastern Asia and North America in the Adiantum pedatum complex (Pteridaceae)

Biogeographic analyses of ferns with an eastern Asian-North American disjunction are few. The Adiantum pedatum complex has such a disjunct distribution. The monophyly of the complex needs to be tested and diversification history of the four species needs to be reconstructed. Plastid (atpA, atpB, rbcL, trnL-F, and rps4-trnS) sequences of 100 accessions representing the biogeographic diversity of Adiantum were analyzed with parsimony and Bayesian inference. Biogeography of the Adiantum pedatum complex was inferred using programs DIVA and LAGRANGE. Divergence times of clades were estimated with the program BEAST. The A. pedatum complex is monophyletic and sister to the eastern Asian A. edentulum. Accessions of A. pedatum do not form a clade; instead three subgroups are recognizable. The clade of A. aleuticum and A. viridimontanum is nested within A. pedatum. The Asian A. myriosorum is sister to the A. pedatum-A. aleuticum clade. Both DIVA and LAGRANGE analyses suggest an eastern Asian origin of the A. pedatum complex. The age of the crown A. pedatum complex is dated to be at 4.27 (2.24-6.57) million years ago. The currently recognized eastern Asian-North American disjunct species A. pedatum needs to be segregated into three species, corresponding to populations in eastern North America, China, and Japan. The eastern Asian-North American disjunction in the complex is inferred to be the result of two intercontinental migrations, one from eastern Asia into North America in the late Tertiary and the other from North America back to eastern Asia in the Pleistocene.

Two more additions to Indian Cololejeunea (Hepaticae: Lejeuneaceae) from West Sikkim

Two species of Cololejeunea (Spruce) Schiffn., viz. C. macounii (Spruce ex Underw.) A. Evans, an Asian-North American disjunct, and C. ocellata (Horik.) Benedix, an East Asian species, are described for the first time in Indian bryoflora from West district of Sikkim.

Molecular phylogeny and biogeographic diversification of Parthenocissus (Vitaceae) disjunct between Asia and North America

Parthenocissus is a genus of the grape family Vitaceae and has a disjunct distribution in Asia and North America with members in both tropical and temperate regions. The monophyly of Parthenocissus has not yet been tested, and the species relationships and the evolution of its intercontinental disjunction have not been investigated with extensive sampling and molecular phylogenetic methods. • Plastid (trnL-F, rps16, and atpB-rbcL) and nuclear GAI1 sequences of 56 accessions representing all 12 Parthenocissus species were analyzed with parsimony, likelihood, and Bayesian inference. Divergence times of disjunct lineages were estimated with relaxed Bayesian dating. Evolution of the leaflet number was assessed by tracing this character onto Bayesian trees using the Trace Character Over Trees option in the program Mesquite. • Parthenocissus is monophyletic and sister to the newly described segregate genus Yua. Two major clades within Parthenocissus are recognizable corresponding to their distribution in Asia and North America. The disjunction between the two continents is estimated to be at 21.64 (95% higher posterior densities 10.23-34.89) million years ago. • Parthenocissus is likely to have derived from the Eocene boreotropical element. Its current Asian-North American disjunction is dated to the early Miocene, congruent with fossil and paleoclimatic evidence. The tropical species is nested within the temperate clade and is inferred to have dispersed from the adjacent temperate regions. Parthenocissus and Yua are best treated as distinct genera. Leaflet number in this genus has a complex history and cannot be used as a character for infrageneric classification.

Palynological insights of the eastern Asian and eastern North American disjunct genus Symplocarpus (Araceae)

Symplocarpus L. (Araceae) is a disjunct genus including S. foetidus (L.) Nutt. var. latissimus (Makino) S. Hara, S. nipponicus Makino, and S. nabekuraensis Otsuka & K. Inoue in temperate eastern Asia; S. egorovii N. S. Pavlova & V. A. Nechaev in the Russian far east; and S. foetidus (L.) Nutt. in temperate northeastern North America. For the first time, pollen morphology of Symplocarpus was investigated by light and scanning electron microscopes. Symplocarpus foetidus var. latissimus and S. foetidus had reticulate surfaces and rounded (obtuse), long, equatorial axis tips, whereas S. nipponicus had microreticulate surfaces and acute tips. Symplocarpus foetidus in North America had larger pollen grains than S. foetidus var. latissimus in eastern Asia. Thus, based on pollen characteristics, S. foetidus var. latissimus is more closely related to S. foetidus than to S. nipponicus. Other lines of evidence, such as molecular phylogenetic studies and phenology, support the phylogenetic relationships among these species proposed in this study. The slightly modified acetolysis procedure conducted in this paper is a promising application to study weak exine or exineless pollen grains.

Development of microsatellite markers for the mossPtychomitrium gardneri(Ptychomitriaceae)

Microsatellite markers were developed for Ptychomitrium gardneri to study population genetics of this eastern Asian-North American disjunct moss. • A total of 13 microsatellite markers were developed in Chinese populations of P. gardneri, using the Fast Isolation by AFLP of Sequences COntaining Repeats protocol. Eight of the markers showed polymorphism when assessed in a sample of four populations of 29 individuals from China. These markers amplified three to four alleles per locus. Five primers also amplified in P. linearifolium and P. wilsonii. • These markers may be useful for further investigation of population genetics of P. gardneri.

Phylogeny and biogeography of the eastern Asian–North American disjunct wild-rice genus ( Zizania L., Poaceae)

The wild-rice genus Zizania includes four species disjunctly distributed in eastern Asia and North America, with three species ( Z. aquatica, Z. palustris, and Z. texana) in North America and one ( Z. latifolia) in eastern Asia. The phylogeny of Zizania was constructed using sequences of seven DNA fragments ( atpB-rbcL, matK, rps16, trnL-F, trnH-psbA, nad1, and Adh1a) from chloroplast, mitochondrial, and nuclear genomes. Zizania is shown to be monophyletic with the North American species forming a clade and the eastern Asian Z. latifolia sister to the North American clade. The divergence between the eastern Asian Z. latifolia and the North American clade was dated to be 3.74 (95% HPD: 1.04–7.23) million years ago (mya) using the Bayesian dating method with the combined atpB-rbcL, matK, rps16, trnL-F, and nad1 data. Biogeographic analyses using a likelihood method suggest the North American origin of Zizania and its migration into eastern Asia via the Bering land bridge. Among the three North American species, the organellar data and the haplotype network of the nuclear Adh1a gene show a close relationship between Z. palustris and the narrowly distributed endangered species Z. texana. Bayesian dating estimated the divergence of North American Zizania to be 0.71 (95% HPD: 0.12–1.54) mya in the Pleistocene. The non-monophyly of Z. palustris and Z. aquatica in the organellar and nuclear data is most likely caused by incomplete lineage sorting, yet low-frequency unidirectional introgression of Z. palustris into Z. aquatica is present in the nuclear data as well.

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.

Systematics and phytogeography of selected Eocene Okanagan Highlands plants

The diverse Early to Middle Eocene Okanagan Highlands floras of south central British Columbia and northeastern Washington reflect a time of rapid evolution and the early radiation of many dicot families that are currently significant elements of temperate floras. Recent studies of the Republic, Washington flora (Klondike Mountain Formation) and related Okanagan floras in British Columbia have documented both the earliest, and sometimes the only, known fossil occurrences of genera. Today many once more widespread taxa are restricted, particularly to Asian and (or) eastern North American refugia. Examples include members of the families Betulaceae (birch, hazelnut), Rosaceae (rose), Hamamelidaceae (witch hazel), and the endemic Asian family Trochodendraceae. Earliest occurrences are noted for Neviusia (Rosaceae), Trochodendron (Trochodendraceae), Corylus and Carpinus (both Betulaceae). The first unequivocal leaf records of Corylopsis and Fothergilla (both Hamamelidaceae), and two new Eocene species of the extinct fruit Palaeocarpinus (Betulaceae) are also recognized. Today, Trochodendron and Corylopsis are restricted to Asia, whereas Neviusia and Fothergilla, genera with close Asian relatives, occur only in North America. Corylus johnsonii from Republic is most similar to the extant Asian species C. heterophylla, C. wangii, and C. ferox. Neviusia leaves from One Mile Creek near Princeton, British Columbia are more similar to N. cliftonii, an endemic from Mount Shasta, California, than to N. alabamensis of southeastern North America. A better documentation of the Okanagan Highlands floras is essential to our understanding of the evolution of North American temperate floras and the nature of Asian – North American disjunct taxa.

Cladistic biogeography of Gleditsia (Leguminosae) based on ndhF and rpl16 chloroplast gene sequences

We used cladistic analysis of chloroplast gene sequences (ndhF and rpl16) to test biogeographic hypotheses in the woody genus Gleditsia. Previous morphological comparisons suggested the presence of two eastern Asian-eastern North American species pairs among the 13 known species, as well as other intra- and inter-continental disjunctions. Results from phylogenetic analyses, interpreted in light of the amount of sequence divergence observed, led to the following conclusions. First, there is a fundamental division of the genus into three clades, only one of which contains both Asian and North American species. Second, the widespread and polymorphic Asian species, G. japonica, is sister to the two North American species, G. triacanthos and G. aquatica, which themselves are closely related inter se, but are both polymorphic and paraphyletic. Third, the lone South American Gleditsia species, G. amorphoides, forms a clade with two eastern Asian species. Gleditsia thus appears to have only one Asian-North American disjunction and no intercontinental species pairs. Low sequence divergence between G. amorphoides and its closest Asian relatives implicates long-distance dispersal in the origin of this unusual disjunction. Sequence divergence between Asian and North American Gleditsia is much lower than between Asian and North American species of its closest relative, Gymnocladus. Estimates of Asian-North American divergence times for Gymnocladus are in general accordance with fossil data, but estimates for Gleditsia suggest recent divergences that conflict with ages of known North American Gleditsia fossils.