Phylogeny and biogeography of Chinese Heracleum (Apiaceae tribe Tordylieae) with comments on their fruit morphology

The Sino-Himalayan Mountain Region is a major center of diversity of Heracleum (Apiaceae, Tordylieae, Tordyliinae), yet the phylogenetic relationships among its rich diversity of endemic taxa are largely unknown. In this study, we (1) investigate the monophyly of Heracleum and ascertain the phylogenetic placements of those species of Heracleum from China that fall outside of the group; (2) estimate phylogenetic relationships within Heracleum sensu stricto, particularly among those species occurring in the Hengduan Mountain Region of southwestern China; and (3) reconstruct the biogeographic history of Heracleum sensu stricto. To achieve these objectives, Bayesian inference and maximum-parsimony analyses of the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) region and five noncoding chloroplast DNA (cpDNA) loci (rps16-trnK, rps16 intron, trnQ-rps16, rpl32-trnL, and psbA-trnH), separately and in combination, were carried out. Fruit morphology of the Chinese Heracleum species was examined. Statistical dispersal-vicariance analysis (S-DIVA) was used to reconstruct the biogeographic history. In total, 114 accessions were considered, which included 88 species representing eight genera of subtribe Tordyliinae, 28 of the 29 species of Heracleum occurring in China, and 4 species of Angelica as outgroups. The genus Heracleum is monophyletic upon the exclusion of nine species of Heracleum from China and the inclusion of the Caucasian monotypic genera Mandenovia and Symphyoloma. These nine excluded species show affinities to Tetrataenium, Semenovia, and Angelica. The species of Heracleum endemic to southwestern China form a moderately to strongly supported clade; however, major discordance between the ITS- and cpDNA-derived phylogenies, lack of resolution in the trees, and polyphyly of several species preclude unequivocal hypotheses of phylogenetic relationship. The distinctly clavate vittae, shorter than the length of the mericarp, is a diagnostic feature for Heracleum sensu stricto. Heracleum originated in the West Palearctic, with three possible migration routes leading to its present-day distribution. Major topological conflicts between the ITS and cpDNA trees may be due to interspecific hybridization and/or incomplete lineage sorting. The taxonomic limits of many Chinese Heracleum species remain unclear, possibly because of recent radiation in the Hengduan Mountains.     

东亚和北美当归属（广义）的果实解剖和演化

应用比较解剖研究了广义当归属在世界主要分布区－东亚和北美种类的果实结构的特征。根据分生果背腹压扁的程度、棱翅与槽的形状、果皮层次和结构、油管数目、大小和位置等稳定性状,可将果实分为５个类型,与《中国植物志》将其分为５个属,即古当归属,当归属,高山芹属,柳叶芹属和山芹属的意见相一致。古当归属为原始的类型,山芹属具有高度演化的特征。且对一些种类有分类位置提出了调整的意见和各属的演化趋势。     

Colchicine-binding protein from phloem and xylem of a higher plant

Summary Colchicine has been demonstrated by ion-exchange and by gelfiltration assay to bind to a protein fraction derived from the higher plant Heracleum mantegazzianum. Colchicine-binding protein from a plant source was much more unstable than tubulin from animal preparations. The tissues of Heracleum vary in their content of colchicine-binding activity. No activity was obtained from non-vascular tissue. Phloem has at most, twice as much activity as xylem. The significance of these results is discussed in relation to a proposed degree of homology between P protein of phloem and microtubule protein.     

Ostericum atropurpureum sp. nov. (Apiaceae) from Zhejiang,China

Ostericum atropurpureum G. Y. Li, G. H. Xia & W. Y. Xie (Apiaceae, Apioideae) from Zhejiang, China, is described and illustrated. It is closely related to O. huadongense Z. H. Pan & X. H. Li and O. sieboldii (Miquel) Nakai, but differs in having leaves with 1.5–9.0 cm long petiole, linear bracteoles 6–12 mm long, 5–9 rays, 7–14‐flowered umbellules, dark purple petals, broadly winged dorsal and lateral fruit ribs, 1.0–1.5 mm broad, 3–6 vittae in each furrow and 4–8 on the commissure.     

湖南珍稀特有植物喜雨草的重新发现及补充描述

喜雨草,湖南珍稀特有植物,属唇形科单型属喜雨草属,自1918年8月韩马迪在湖南西南部武冈云山采到模式标本,并于1936年作为一个新属、新种发表后,其踪迹消失全无。该次报道为喜雨草被发现近100年后,在湖南宁远、通道、道县、新宁被重新发现。根据采集的标本和拍摄的照片,更正修改和补充了原形态描述,重新确定了该种花萼为3/2式,花冠颜色为白色。     

Karyotype and cytogeography of the genus Heracleum (Apiaceae) in the Hengduan Mountains

Abstract In the present study, the karyotypes of 34 populations belonging to 11 species and one variety of Heracleum from the Hengduan Mountains in China were examined. Chromosome numbers and the karyotypes of three species (H. souliei, H. kingdoni, and H. wenchuanense) are reported for the first time, as are the karyotypes of H. moellendorffii and H. henryi (tetraploid). Populations of H. candicans, H. franchetii, and H. kingdoni in the Hengduan Mountains were found to consist of a mixture of diploid and tetraploid plants. Except for four species of Heracleum, namely H. candicans, H. franchetii, H. henryi, and H. kingdoni, which have both diploid and tetraploid karyotypes, all other species of Heracleum are were found to be diploid. All karyotypes were found to belong to the 2A type of Stebbins, with the exception of H. candicans var. obtusifolium, which belongs to 2B, and H. hemsleyanum and H. franchetii (Mt. Dujuan, Daocheng, Sichuan, China), which belong to 1A. There was only a slight difference in the karyotype asymmetry index, which suggests a close kinship for species of Heracleum and that the entire phylogenetic development of Heracleum is relatively primitive. Species that exhibited advanced morphological features were also more advanced in karyotype structure, with the order of karyotype evolution being 1A→2A→2B. This phenomenon indicates that the species distributed in the Hengduan Mountains have not diverged completely and that the Hengduan Mountains are a relatively young and active area for the evolution of Heracleum. Polyploidization in Heracleum may be an important evolutionary mechanisms for some species, generating diversity. The biological attributes, distribution range, and the geological history of the genus have all played a part in accelerating the evolution through polyploidization or aneuploidization. It is known that as the distribution latitude of Heracleum decreases from north to south, the chromosome number, ploidy level, and asymmetry structure appear to increase. In the Hengduan Mountains, these tendencies are also evident. Finally, based on all the available cytogeographic data, we speculate that the more advanced tetraplont or aneuploid species of Heracleum in India may be derived from early diplont species that were distributed in the Caucasus region and Hengduan Mountains. The dispersal of Heracleum was from Eurasia to India, because this correlates with the emergence of the Himalayan Mountains through tectonic movement. Thus, the Hengduan Mountains are not only a center of diversity for Heracleum, but also a center of active speciation in modern times.     

Invasion by Heracleum mantegazzianum in different habitats in the Czech Republic

Abstract. Heracleum mantegazzianum, a tall forb from the western Caucasus invaded several different habitats in the Czech Republic. The relation between invasion success and type of recipient habitat was studied in the Slavkovskù les hilly ridge, Czech Republic. The vegetation of 14 habitat types occurring in an area of ca. 25 km² was analysed using phytosociological relevés, and the invasion success of Heracleum (in terms of number of localities, area covered and proportion of available area occupied) was recorded separately in each of them. Site conditions were expressed indirectly using Ellenberg indicator values. The hypothesis tested was that Heracleum spreads in the majority of vegetation types regardless of the properties of the recipient vegetation. Community invasibility appeared to be affected by site conditions and the composition of the recipient vegetation. The species is not found in acidic habitats. Disturbed habitats with good possibilities of dispersal for Heracleum seeds are more easily invaded. Communities with a higher proportion of phanerophytes and of species with CS (Competitive/Stresstolerating) strategy were more resistant to invasion. The invasion success was bigger in sites with increased possibilities of spread for Heracleum diaspores. Communities invaded by Heracleum had a lower species diversity and a higher indicator value for nitrogen than not-invaded stands. It appears that species contributing to community resistance against invasion of Heracleum, or capable of persisting in Heracleum-invaded stands, have similar ecological requirements but a different life strategy to the invader.     

Gas exchanges of three co-occurring species of Cypripedium in a scrubland in the Hengduan Mountains

Gas exchanges and related leaf traits of three co-occurring species of genus Cypripedium (C. yunnanense Franch., C. guttatum SW., and C. flavum P.F. Hunt et Summerch.) were investigated in a scrubland at 3 460 m a.s.l. in the Hengduan Mountains. The considered species had similar photosynthetic responses to photosynthetic photon flux density (PPFD) and air temperature. The photosynthetic capacity (P _max), carboxylation efficiency (CE), apparent quantum efficiency (AQE), PPFD-saturated rate of electron transport (J_max), respiration rate (R _D), and leaf nitrogen content per unit area (LNC) of C. guttatum were higher than those of C. yunnanense and C. flavum. The highest P _max of C. guttatum was related to the highest LNC and the lowest ratio of intercellular CO₂ concentration to atmospheric CO₂ concentration (C _i/C _a). However, no significant differences in stomatal conductance (g _s) and relative stomatal limitations (RSL) were observed among the three species. Hence biochemical limitation had a dominant role in P _max differences among the considered species.     

高分辨裂解—气相色谱在前胡族系统分类中的应用

采用高分辨裂解－气相色谱法对前胡族（Peucedaneae Drude)20属35种植物的叶片成分进行分析,经过聚类分析得出以下结果：35种植物基本可分为3大类,对应于3个亚族。阿魏亚族（Ferulinae Drude)的球根阿魏属（Schumannia Kuntze)、伊犁芹属（Talassia Korov.)和簇花芹属（Soranthus Ledeb.)均应为独立属。环翅芹亚族（Tordyliinae Drude)的四带芹属[Tetrataenium(DC.)Manden.]和大瓣芹属（Semenovia Regel et Herd.)二者也作为独立属处理为宜,其中有争议的锐尖叶独活[Heracleum longilobum(Norman)Sheh et T.S.Wang]不应作为大瓣芹属成员,仍应保留于独活属。当归亚族（Angelicinae Drude)的山芎[Conioselinum chinense(L.)Britton]和东当归[Angelica acutiloba(Sieb.et Zucc.)Kitagawa]曾被作为阿米芹族（Ammineae Koch)的蛇床属（Cnidium Cuss.)或藁本属（Ligusticum L.)的成员,聚类图显示其属于前胡族的当归亚族是合适的。     

ITS phylogeny of West Asian Heracleum species and related taxa of Umbelliferae–Tordylieae W.D.J.Koch, with notes on evolution of their psbA-trnH sequences

Summary Heracleum is a large and taxonomically complex genus of the Umbelliferae–Tordylieae. The phylogenetic relationships of West Asian Heracleum species and related taxa were explored using data from sequences of the internal transcribed spacer (ITS1 and ITS2) region of nuclear ribosomal DNA. The data set consists of 56 species, of which 47 were analyzed for the first time; it represents all subdivisions of the genus Heracleum, as well as some representatives of Pastinaca complex. Heracleum was shown to be a polyphyletic genus, as its species fall into two different clades, one of which comprises also Symphyoloma and Mandenovia. Section Pubescentia was confirmed, in contrast to the sections Villosa and Heracleum being polyphyletic. A separate position of the section Wendia was supported. H. marashicum was shown to be a member of a clade comprising Pastinaca and related genera. The sequences of chloroplast psbA-trnH intergenic spacer, the region recently proposed for DNA barcoding in plants, were also analyzed for 33 species, representing all principal clades within Heracleum and its relatives. They have been proven to be very similar and not suitable for DNA barcoding in this group. However, some sequence variation was revealed. This variation could be explained by the combination of such evolutionary events as inversion and duplication. It was shown that these events are rather common in Tordylieae and can occur independently in different lineages. The evolution patterns of psbA-trnH spacer are hypothesized.     

The teleomorph of Leptographium yunnanense, discovered in crosses among isolates from Thailand, China, and Japan

A teleomorph was discovered in crosses among isolates of Leptographium yunnanense isolated from Pinus spp. originating in Thailand, China, and Japan. The ascocarps are black, globose to subglobose, and lacking necks. Ascospores are hyaline, 1-celled, surrounded by hyaline sheaths, and appear cucullate in side view, quadrangular in face view, and triangular in end view. Three species were known to have teleomorphs morphologically similar to the present fungus. However, their anamorphs were distinguishable from L. yunnanense. Thus, this teleomorph is described as Grosmannia yunnanensis. Contribution No. 214, Laboratory of Plant Parasitic Mycology, Graduate School of Life & Environmental Sciences, University of Tsukuba.     

über den Feinbau von öldrüsen

Zusammenfassung Die ölkanÄle vonHeracleum sphondylium undDorema ammoniacum wurden elektronen-mikroskopisch untersucht, um Aufschlüsse über den Exkretionsproze\ zu gewinnen. Die Drüsenepithelzellen vonHeracleum enthalten zahlreiche glatte, tubulÄre Elemente des endoplasmatischen Reticulums. Die Leukoplasten sind teilweise vom endoplasmatischen Reticulum eingehüllt. Sie enthalten viele tubulÄre Strukturen. Die Dictyosomen bilden Sekretvesikel, die vermutlich Wandmaterial und Schleim extruieren. Die Drüsenzellen vonDorema sind im Prinzip Ähnlich gebaut. Jedoch haben die Leukoplasten noch mehr Tubuli, und das tubulÄre, glatte endoplasmatische Reticulum ist nicht so stark entwickelt. Der Kanalinhalt ist anders zusammengesetzt. Die Ergebnisse werden mit Angaben aus der Literatur verglichen, und verschiedene Möglichkeiten für den Weg des Exkretes werden diskutiert.

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The fine structure of oil glandsIV. The oil channels of umbelliferae:Heracleum sphondylium andDorema ammoniacum

Summary The oil channels ofHeracleum sphondylium andDorema ammoniacum were studied by electron microscopy in different stages of development with regard to the excretion process. Characteristic features of theHeracleum gland cells are a smooth endoplasmic reticulum consisting of a network of tubules, elements of the endoplasmic reticulum which envelope partly the leucoplasts, and irregular tubular structures within these plastids. The dictyosomes extrude secretion vesicles which may contain wall substances and the mucilagous constituents of the excretion. The content of the channel is poorly preserved in its arrangement. The gland cells ofDorema have a similar structure. There are certain differences in the content of the channel. The tubular elements within the leucoplasts are more prominent. The smooth, tubular endoplasmic reticulum is not so conspicuous. The results were compared with other reports, and possibilities for the way of the excretion were discussed.

     

Bupleurum shanianum sp. nov. (Apiaceae) from China

Bupleurum shanianum X. G. Ma & X. J. He, previously recognized in China as B. gracillimum, is proposed here as a new species. This new species and B. gracillimum differ in many morphological characters (flowering stem position, bract size, number of flowers per umbellule) and have non‐overlapping distribution ranges. Bupleurum shanianum is most similar to B. yunnanense Franch., but the new species has shorter cauline leaves, and umbellules usually with 4 bracteoles and 3–6(–9) flowers. Bupleurum shanianum is known from eastern Himalayas to western Sichuan.     

Pternopetalum arunachalense sp. nov. (Apiaceae) from India

Pternopetalum arunachalense M. Bhaumik & P. Satyanar. is a new species from Arunachal Pradesh in India and is described and illustrated here. This new species is similar to P. trichomanifolium (Franch.) Hand.‐Mazz. by highly dissected, ternate 2–4‐pinnate leaves, ultimate segments linear and erect elongate styles but differs by generally smaller habit and scabrid mericarps with 1–2 vittae in each furrow and commissure.     

Sinephropteris,a new genus of scolopendrioid ferns

A new genus of ferns,Sinephropteris, is described from southeast Asia. Its one species,S. delavayi (Franch.) Mickel, has most recently been placed inSchaffneria, an otherwise monotypic Mexican genus, but there is evidence that they are not closely related.     

9种日本当归属植物的果实形态特征研究

当归属是伞形科的代表性大属,日本列岛是其重要的分布地和多样性中心之一。该研究对产自日本的9种当归属植物(特有种8种)的果实形态结构和表面微形态特征进行了观察和比较,以明确它们的分类学特征,探讨潜在的种间关系,为进一步的分类修订和系统发育研究提供资料和依据。结果表明:(1)体式显微镜观察发现,产自日本的9种当归属植物的果实形状主要为倒卵形和长圆形,表面光滑或少毛,呈不同程度的背腹压扁,背棱大多呈线状突起或狭翅状,侧棱则呈或宽或窄的翅状,油管一般为每个棱槽1条、合生面2～4条。(2)扫描电镜观察发现,大多数材料的果实具有疏密不等的平行线形初级蜡质纹饰,次级纹饰呈现鳞状、束状、索状、丘状或不明显,个别可见珊瑚状或网格状基底,一般具有颗粒状附属物。(3)通过分析比较上述形态特征和鉴别要点,编制了分种检索表,并结合中国当归属系统发育的研究成果,进一步对日本特有当归属植物的系统位置及其与东亚大陆物种的亲缘关系进行了讨论。研究认为,日本特有当归属植物的果实形态特征较丰富,能为种间分类和亲缘关系研究提供有力支持。     

Invasive species of Heracleum in Europe: an insight into genetic relationships and invasion history

Several species of the genus Heracleum (Umbelliferae) were introduced into Europe from south-west Asia in the 19th century and are now widespread in many countries. At least three invasive taxa with unresolved relationships to one another are thought to occur in Europe: Heracleum mantegazzianum Sommier & Levier, H. sosnowskyi Manden, and H. persicum Desf. ex Fischer. They are tall plants forming extensive stands with a high cover. To elucidate genetic relationships between the species, and gain insight into their invasion history, samples were collected from native ranges in Asia and invaded ranges of the three species in Europe and analysed using amplified fragment length polymorphism. Five other Heracleum species were also studied and in total, 189 samples from 72 populations were analysed. The results confirmed that there are three distinct tall Heracleum species invading in Europe. Within each of the three species, plants collected in the invaded range are genetically close to those from their native ranges. A close genetic relationship between the three invasive Heracleum species in Europe was also found. A high overall genetic variability detected in the invaded range suggests that the majority of invading populations were not affected by a genetic bottleneck and that rapid evolution, drift, or hybridization played a role in genetic structuring of invading populations. For H. mantegazzianum , genetic distance of populations in the native range significantly decreased with geographical distance, but not in the invaded range. It is likely that the current pattern of genetic diversity in Europe resulted from multiple introductions of all three species.     

Billburttia, a new genus of Apiaceae (tribe Apieae) endemic to Madagascar

The genus Billburttia is described to include two new species endemic to Madagascar, B. capensoides and B. vaginoides. Both species were tentatively placed within the problematic genus Peucedanum L. (as Peucedanum sp. A and Peucedanum sp. B) based on their dorsally compressed fruits lacking prominent dorsal ribs and with winged marginal ribs. Recently, however, the African members of Peucedanum have been shown to be only distantly related to the type of the genus and have therefore been segregated into six African endemic genera. While the Malagasy species appear superficially similar to members from one of these segregate genera, viz. Notobubon, they differ markedly in their fruit anatomical characters, notably a narrower commissure, six commissural vittae, vascular tissue in the tip of the ribs and sphaerocrystals distributed in and around the epidermis. The latter two characters apparently represent apomorphies for Billburttia. The non-peucedanoid affinity of the genus, as suggested by the fruit anatomical data, was confirmed using ITS and rps16 intron sequences. Both parsimony and Bayesian analyses of these data place Billburttia within the tribe Apieae of subfamily Apioideae and not closely related to either Peucedanum (Selineae) or the African peucedanoid genera (Lefebvrea clade of Tordylieae).