A monograph of Caprifoliaceae: Linnaeeae

Tribe Linnaeeae is a diverse group disjunctly distributed in the northern hemisphere between China and Japan and Mexico. The tribe consists of shrubs with showy flowers that are important in the horticultural industry. We present a detailed monograph of tribe Linnaeeae that includes descriptions, identification keys, botanical plates with line drawings and photographs, distribution maps, morphometric analysis of key characters, and discussion on evolution, cultivation and conservation. Twenty species, as well as five new varieties and four new forms, are recognised in six genera. Forty heterotypic names (not including the 218 names for Linnaea borealis) are listed as synonyms. 1,973 specimens were studied and databased from 13 herbaria (A, BM, CAS, E, GZAC, IBSC, KUN, KYO, MAK, PE, TNS, TOFO and TI). The Kolkwitzia–Dipelta–Diabelia clade radiated from the Qinling Mountain Range of China and is disjunctly distributed in Japan. The Linnaea–Vesalea clade may have migrated through the Bering Land Bridge and radiated along the Sierra Madre Oriental in Mexico. Abelia, which is most diverse in China, evolved in relation to the formation of the Qinghai Tibetan Plateau

Complete chloroplast genome of Nyctocalos pinnatum: chloroplast features and phylogenetic relationships within Bignoniaceae

Nyctocalos is a genus of flowering lianas belonging to the family Bignoniaceae, and occurring from South-Central China to Malesia. In this study, we assembled the first complete chloroplast genome of N. pinnatum. The total length of the chloroplast genome is 159591 bp, with a GC content of 38.04%, which includes a pair of inverted repeats of 30,480 bp, a small single-copy region of 12,774 bp and a large single-copy region of 85,857 bp. The chloroplast genome contains 135 genes, consisting of 89 protein-coding genes, 38 transfer RNAs, and 8 ribosomal RNAs. We constructed a phylogenomic tree with representative chloroplast genomes from Bignoniaceae. N. pinnatum is revealed to be sister to Oroxylum in the tribe Oroxyleae, with a high bootstrap support. This is the first chloroplast genome assembled in Nyctocalos, and it provides essential information for further ecology and evolutionary studies in this genus and Bignoniaceae

Plastid phylogenomic insights into the evolution of the Caprifoliaceae s.l. (Dipsacales)

The family Caprifoliaceae s.l. is an asterid angiosperm clade of ca. 960 species, most of which are distributed in temperate regions of the northern hemisphere. Recent studies show that the family comprises seven major clades: Linnaeoideae, Zabelia, Morinoideae, Dipsacoideae, Valerianoideae, Caprifolioideae, and Diervilloideae. However, its phylogeny at the subfamily or genus level remains controversial, and the backbone relationships among subfamilies are incompletely resolved. In this study, we utilized complete plastome sequencing to resolve the relationships among the subfamilies of the Caprifoliaceae s.l. and clarify several long-standing controversies. We generated and analyzed plastomes of 48 accessions of Caprifoliaceae s.l., representing 44 species, six subfamilies and one genus. Combined with available Caprifoliaceae s.l. plastomes on GenBank and 12 outgroups, we analyzed a final dataset of 68 accessions. Genome structure was strongly conserved in general, although the boundaries between the Inverted Repeat were found to have contracted across Caprifoliaceae s.l. to exclude rpl2, rps19, and ycf1, all or parts of which are typically present in the IR of most angiosperms. The ndhF gene was found to have been inverted in all plastomes of Adoxaceae. Phylogenomic analyses of 68 complete plastomes yielded a highly supported topology that strongly supported the monophyly of Zabelia and its sister relationship to Morinoideae. Moreover, a clade of Valerianoideae + Dipsacoideae was recovered as sister to a clade of Linnaeoideae + Zabelia + Morinoideae clade, and Heptacodium was sister to remaining Caprifolioideae. The Diervilloideae and Caprifolioideae were successively sister to all other Caprifoliaceae s.l. Major lineages of Caprifoliaceae s.l. were estimated to have diverged from the Upper Cretaceous to the Eocene (50-100 Ma), whereas within-genus diversification was dated to the Oligocene and later, concomitant with global cooling and drying. Our results demonstrate the power of plastid phylogenomics in improving estimates of phylogeny among genera and subfamilies, and provide new insights into plastome evolution across Caprifoliaceae s.l

Plastid phylogenomic insights into the evolution of the Caprifoliaceae \u3ci\u3es.l.\u3c/i\u3e (Dipsacales)

The family Caprifoliaceae s.l. is an asterid angiosperm clade of ca. 960 species, most of which are distributed in temperate regions of the northern hemisphere. Recent studies show that the family comprises seven major clades: Linnaeoideae, Zabelia, Morinoideae, Dipsacoideae, Valerianoideae, Caprifolioideae, and Diervilloideae. However, its phylogeny at the subfamily or genus level remains controversial, and the backbone relationships among subfamilies are incompletely resolved. In this study, we utilized complete plastome sequencing to resolve the relationships among the subfamilies of the Caprifoliaceae s.l. and clarify several long-standing controversies. We generated and analyzed plastomes of 48 accessions of Caprifoliaceae s.l., representing 44 species, six subfamilies and one genus. Combined with available Caprifoliaceae s.l. plastomes on GenBank and 12 outgroups, we analyzed a final dataset of 68 accessions. Genome structure was strongly conserved in general, although the boundaries between the Inverted Repeat were found to have contracted across Caprifoliaceae s.l. to exclude rpl2, rps19, and ycf1, all or parts of which are typically present in the IR of most angiosperms. The ndhF gene was found to have been inverted in all plastomes of Adoxaceae. Phylogenomic analyses of 68 complete plastomes yielded a highly supported topology that strongly supported the monophyly of Zabelia and its sister relationship to Morinoideae. Moreover, a clade of Valerianoideae + Dipsacoideae was recovered as sister to a clade of Linnaeoideae + Zabelia + Morinoideae clade, and Heptacodium was sister to remaining Caprifolioideae. The Diervilloideae and Caprifolioideae were successively sister to all other Caprifoliaceae s.l. Major lineages of Caprifoliaceae s.l. were estimated to have diverged from the Upper Cretaceous to the Eocene (50–100 Ma), whereas within-genus diversification was dated to the Oligocene and later, concomitant with global cooling and drying. Our results demonstrate the power of plastid phylogenomics in improving estimates of phylogeny among genera and subfamilies, and provide new insights into plastome evolution across Caprifoliaceae s.l

Molecular Phylogeny and Biogeographic Diversification of Linnaeoideae (Caprifoliaceae s. l.) Disjunctly Distributed in Eurasia, North America and Mexico

Linnaeoideae is a small subfamily of erect or creeping shrubs to small trees in Caprifoliaceae that exhibits a wide disjunct distribution in Eurasia, North America and Mexico. Most taxa of the subfamily occur in eastern Asia and Mexico but the monospecific genus Linnaea has a circumboreal to north temperate distribution. In this study, we conducted phylogenetic and biogeographic analyses for Linnaeoideae and its close relatives based on sequences of the nuclear ribosomal ITS and nine plastid (rbcL, trnS-G, matK, trnL-F, ndhA, trnD-psbM, petB-D, trnL-rpl32 and trnH-psbA) markers. Our results support that Linnaeoideae is monophyletic, consisting of four eastern Asian lineages (Abelia, Diabelia, Dipelta and Kolkwitzia), the Mexican Vesalea, and Linnaea. The Mexican Vesalea was formerly placed in Abelia, but it did not form a clade with the eastern Asian Abelia; instead Vesalea and Linnaea are sisters. The divergence time between the eastern Asian lineages and the Mexican Vesalea plus the Linnaea clade was dated to be 50.86 Ma, with a 95% highest posterior density of 42.8 Ma (middle Eocene) to 60.19 Ma (early Paleocene) using the Bayesian relaxed clock estimation. Reconstructed ancestral areas indicated that the common ancestor of Linnaea plus Vesalea may have been widespread in eastern Asia and Mexico or originated in eastern Asia during the Eocene and likely migrated across continents in the Northern Hemisphere via the North Atlantic Land Bridges or the Bering Land Bridge. The Qinling Mountains of eastern Asia are the modern-day center of diversity of Kolkwitzia-Dipelta-Diabelia clade. The Diabeliaclade became highly diversified in Japan and eastern China. Populations of Diabelia serrata in Japan and eastern China were found to be genetically identical in this study, suggesting a recent disjunction across the East China Sea, following the last glacial event

Molecular Phylogeny and Biogeographic Diversification of Linnaeoideae (Caprifoliaceae s. l.) Disjunctly Distributed in Eurasia, North America and Mexico

Linnaeoideae is a small subfamily of erect or creeping shrubs to small trees in Caprifoliaceae that exhibits a wide disjunct distribution in Eurasia, North America and Mexico. Most taxa of the subfamily occur in eastern Asia and Mexico but the monospecific genus Linnaea has a circumboreal to north temperate distribution. In this study, we conducted phylogenetic and biogeographic analyses for Linnaeoideae and its close relatives based on sequences of the nuclear ribosomal ITS and nine plastid (rbcL, trnS-G, matK, trnL-F, ndhA, trnD-psbM, petB-D, trnL-rpl32 and trnH-psbA) markers. Our results support that Linnaeoideae is monophyletic, consisting of four eastern Asian lineages (Abelia, Diabelia, Dipelta and Kolkwitzia), the Mexican Vesalea, and Linnaea. The Mexican Vesalea was formerly placed in Abelia, but it did not form a clade with the eastern Asian Abelia; instead Vesalea and Linnaea are sisters. The divergence time between the eastern Asian lineages and the Mexican Vesalea plus the Linnaea clade was dated to be 50.86 Ma, with a 95% highest posterior density of 42.8 Ma (middle Eocene) to 60.19 Ma (early Paleocene) using the Bayesian relaxed clock estimation. Reconstructed ancestral areas indicated that the common ancestor of Linnaea plus Vesalea may have been widespread in eastern Asia and Mexico or originated in eastern Asia during the Eocene and likely migrated across continents in the Northern Hemisphere via the North Atlantic Land Bridges or the Bering Land Bridge. The Qinling Mountains of eastern Asia are the modern-day center of diversity of Kolkwitzia-DipeltaDiabelia clade. The Diabeliaclade became highly diversified in Japan and eastern China. Populations of Diabelia serrata in Japan and eastern China were found to be genetically identical in this study, suggesting a recent disjunction across the East China Sea, following the last glacial event

Plastome phylogenomic insights into the Sino-Japanese biogeography of Diabelia (Caprifoliaceae)

Understanding the causes of the Sino-Japanese disjunctions in plant taxa has been a central question in eastern Asian biogeography, with vicariance or long-distance dispersal often invoked to explain such patterns. Diabelia Landrein (Caprifoliaceae; Linnaeoideae) comprises four shrubby species with a Sino-Japanese disjunct distribution. The species diversification time within Diabelia, covering a long geological history of the formation process of the Sino-Japanese flora, dated back to the middle Oligocene, therefore, Diabelia would be an ideal model to elucidate the biogeographic patterns of Sino-Japanese disjunctions with climate fluctuation. In this study, we analyzed complete plastome sequence data for 28 individuals representing all four species of Diabelia. These 28 plastomes were found to be highly similar in overall size (156 243-157 578 bp), structure, gene order, and content. Our phylogenomic analysis of the plastomes supported a close relationship between Diabelia ionostachya (Nakai) Landrein & R.L. Barrett var. wenzhouensis (S.L. Zhou ex Landrein) Landrein from eastern China and Diabelia spathulata (Siebold & Zucc.) Landrein var. spathulata from Japan. Diabelia serrata (Siebold & Zucc.) Landrein was identified as sister to a population of Diabelia sanguinea (Makino) Landrein from Tochigi in central Japan and D. spathulata Landrein, from Toyama, central Japan. Most Diabelia lineages were estimated to have differentiated 8-28 Mya. Our results indicate that two independent vicariance events could explain the disjunction between Japan and Korea in the mid to late Miocene, and between Zhejiang and Japan in the early Miocene

Aristolochia tongbiguanensis, a new species of Aristolochiaceae from Yunnan, China

Aristolochia tongbiguanensis (Aristolochiaceae), a new species from Dehong Prefecture, Yunnan, China, is described and illustrated. It is compared with morphologically similar species including A. faviogonzalezii, A. saccata, A. balansae, A. tonkinensis and A. cathcartii. This new species differs mainly by the shape and colour of its calyx. The perianth of A. tongbiguanensis is yellowish-white with dark purple lines, the limb is nearly rectangular and covered with purple warts as well as long papillae and the throat is yellowish-white with dark purple lines and dots. Full description of A. tongbiguanensis is provided, together with line drawings, photographs and a conservation assessment for this rare vine species

Phylogeography and Population History of Eleutharrhena macrocarpa (Tiliacoreae, Menispermaceae) in Southeast Asia’s Most Northerly Rainforests

The diversification of Tiliacoreae and the speciation of Eleutharrhena are closely linked to Southeast Asia’s most northerly rainforests which originate from the Himalayan uplift. Migration routes across biogeographical zones within the Asian clade, including those of Eleutharrhena, Pycnarrhena, and Macrococculus, and their population structures are still unexplored. We combine endocarp morphology, phylogenetic analyses, divergence time estimation, ancestral area reconstruction, as well as SCoT method to reconstruct the past diversification of Eleutharrhena macrocarpa and to understand their current distribution, rarity, and evolutionary distinctiveness. The disjunct, monospecific, and geographically restricted genera Eleutharrhena and Macrococculus both have a dry aril, a unique feature in Menispermaceae endocarps that further confirms their close relationship. Pycnarrhena and Eleutharrhena appeared during the end of the Oligocene c. 23.10 million years ago (Mya) in Indochina. Eleutharrhena speciation may be linked to climate change during this time, when humid forests became restricted to the northern range due to the Himalayan uplift. Differentiation across the Thai–Burmese range could have contributed to the isolation of the Dehong populations during the Miocene c. 15.88 Mya, when exchange between India and continental Asia ceased. Dispersal to the Lanping–Simao block and further differentiation in southeastern and southern Yunnan occurred during the Miocene, c. 6.82 Mya. The specific habitat requirements that led to the biogeographic patterns observed in E. macrocarpa contributed to a low genetic diversity overall. Population 1 from Dehong, 16 from Pu’er, and 20 from Honghe on the East of the Hua line have a higher genetic diversity and differentiation; therefore, we suggest that their conservation be prioritized