A phylogenetic analysis of the Schisandraceae based on morphology and nuclear ribosomal ITS sequences

Cladistic analyses were conducted using morphological and ITS sequence data to examine phylogenetic relationships within the Schisandraceae. The most parsimonious trees obtained from the morphological data consistently show a dichotomy, with clades corresponding with the two currently accepted genera, Kadsura and Schisandra . In the ITS trees S. propinqua consistently groups with the Kadsura clade, and the position of S. glabra is uncertain. The trees resulting from the analysis of combined data are polytomous, due to the variable positions of K. scandens, K. coccinea and S. propinqua . The monophyletic status of Kadsura subgen. Kadsura sect. Kadsura and Schisandra subgen. Pleiostema are strongly supported, although the monophyly of other supraspecific taxa is unclear. Further studies are required before an unambiguous phylogeny is achieved for the family.     

Micromorphological features of the seed surface of Schisandraceae and their systematic significance

Micromorphological features of the seed surface of the Schisandraceae are reported for the first time. One hundred and seventeen seed samples from 92 populations, representing 24 species, seven varieties and one form of the Schisandraceae, were examined and photographed under scanning electron microscope (SEM). Micromorphological features of seed surfaces of the Schisandraceae are little affected by the habitats in which plants grow, and are quite constant within species, therefore they can be used as reliable diagnostic characters to distinguish species. The following taxonomic treatments are not supported by micromorphological features observed: ( 1 ) To reduce Schisandra wilsoniana A. C. Smith to S. henryi Clarke; (2) To reduce S. neglecta A. C. Smith, S. arisanensis Hayata, S. viridis A. C. Smith, S. sphenanthena Rehd. & Wils., S. gracilis A. C. Smith, S. micrantha A. C. Smith and S. lancifolia var. polycarpa Z. He to S. elongata ( Bl. ) Baill.; (3) To reduce S. henryi var. longipes (Merr. & Chun) A. C. Smith, S. tomentella A. C. Smith and S. pubescens var. pubinervis (Rehd. & Wils.)A. C. Smith to S. pubescens Hemsl. & Wils.; (4) To reduce S. rubriflora(Franch.)Rehd. & Wils., S. flaccidiramosa C. R. Sun, S. incarnata Stapf, S. sphaerandra Stapf, S. sphaerandra f. pallida A. C. Smith and S. glaucescens Diels to S. grandiflora ( Wall. ) Hook. f. & Thoms.; (5) To reduce Schisandra wilsoniana to S. bicolor Cheng; (6) To reduce S. lancifolia var. polycarpa to S. neglecta; (7) To raise S. henryi var. longipes to S. longipes(Merr. & Chun)R. M. K. Saunders; (8) To reduce Kadsura polysperma Yang to K. heteroclita(Roxb. )Craib. The relationship between Schisan-dra Michx. and Kadsura Kaempf. ex Juss. revealed by the micromorphological features of seed surface is very similar to that revealed by pollen morphology, namely these two genera might have originated from a common ancestor and then evolved along two different routes. However the probability of a more complicated relationship between the two genera than we have known hitherto can not be excluded. The general evolutionary level of Kadsura seems to be higher than that of Schisandra. Therefore, the viewpoint that Kadsura is more primitive than Schisandra is not supported.     

国产五味子科植物导管分子的比较解剖

对五味子科 2属 1 9种植物导管分子的结构进行了比较观察。结果表明 :2属均以具缘纹孔导管为主 ,少有梯纹和螺纹导管 ,南五味子属多数种还有梯孔纹导管 ;2属导管长度和宽度差异很小 ,长 /宽比值有差异 ;根据穿孔板的结构可分为 2种形式 :( 1 )单穿孔板 ,这类穿孔板较普遍存在 ;( 2 )梯形穿孔板 ,发现 2属中的红花五味子 (Schisandrarubriflora)、五味子 (S .chinensis)、翼梗五味子 (S .henryi)、铁箍散 (S .propinquavar.sinensis)和黑老虎 (Kadsuracoccinea) 5种植物的导管分子具有此类穿孔板。其中五味子属中的五味子导管分子只具有梯形穿孔板 ,无单穿孔板 ,但横条较少 ,多为 2～ 3条。     

五味子科比较木材解剖及其系统学意义

在光学显微镜和扫描电镜下观察了南五味子属Kadsura 7种21个样品和五昧子属Schisandra 8种14个样品的木材解剖特征,结果表明次生木质部的导管分子类型、导管一射线间纹孔的排列方式、射线类型、射线细胞形状等性状在科的水平上很稳定,这些共同特征都支持五味子科Schisandraceae是比较自然的类群。在五昧子科中发现木材导管单生、具梯状穿孔板、导管壁具梯形排列的纹孔以及木射线异型等原始性状,支持五味子科在被子植物中的原始地位。此外,该科木材还具有单穿孔板导管、导管次生壁具螺纹加厚、具分隔纤维等较为特化的性状状态,这种性状进化水平的异等级现象,使五味子科表现出不同进化水平性状的镶嵌组合。根据木材解剖性状对五味子科进行UPGMA聚类分析,所得结果显示南五昧子属和五味子属在木材解剖特征方面有一定的交叉和重叠,这与分子系统学的结论一致,表明这两个属关系密切,可能起源于共同的祖先。通过比较五昧子科与八角科Illiciaceae的木材解剖特征,进一步证明两个科的亲缘关系很近,不支持将五味子科从八角目Illiciales中独立出来成立五味子目Schisandrales的观点。     

五味子科的系统发育:核糖体DNA ITS区序列证据

The Schisandraceae is one of the most important taxa for understanding the origin and evolution of primitive angiosperms due to its basal position in the recent cladograms of the angiosperm based both on several gene sequences and on morphological characters, but phylogenetic relationships within the family are still unresolved. The sequences of nrDNA ITS region of 15 species representing four sections of Schisandra Michx., two sections of Kadsura Kaempf. ex Juss. and one outgroup, Illicium fargesii Finet et Gagnep., were used to reconstruct the phylogeny of Schisandraceae. Fourteen most parsimonious trees (Length=259, CI=0.934 and RI=0.889) were obtained from the analysis with I．fargesii as the outgroup. In the consensus tree, the genus Schisandra was found to be divided into three clades. Sect. Pleiostema formed a clade together with sect. Maximowiczia, sect. Sphaerostema was weakly supported to be the sister group of a clade comprising S. bicolor var. tuberculata and Kadsura species. In particular, S. bicolor var. tuberculata, a species of sect. Schisandra, was nested within Kadsura. It seems from this result and the morphological characters that Schisandra might not be a monophyletic group. According to the present molecular phylogeny, both elongated hypocarpium and deciduous habit originated independently at least twice in the Schisandraceae, and therefore, the value of the present morphological characters used in the classification of the family Schisandraceae should be reevaluated.     

国产五味子科五种植物叶片脉序研究

首次报道了国产五味子科５种植物的叶脉特征,对科、属、种的特征作了描述,编排有分种检索表．通过与八角科叶脉的比较,支持建立五味子科与八角科的观点,认为五味子属的系统位置在南五味子属之后,并讨论了八角目的演化趋势     

五味子科几个分类学问题探讨

基于近年来的研究成果,对五味子科的分类地位和系统关系、五味子属和南五味子属之间的关系及其分类系统等问题进行了讨论。五味子科和八角科不仅亲缘关系较近,而且是现存被子植物中最早分化出来的类群,把它们同置于八角目是合理的。五味子科的两个属——五味子属和南五味子属,可能源于同一祖先并沿两条不同演化路线平行演化。目前,关于南五味子属的分类系统的意见比较一致,但对五味子属分类系统的分歧却很大。     

五味子科植物的木脂素类成分及生物活性与国内资源

五味子料(Schisandraceae)是一类重要的药用植物.本文简述从国产的12种五味子植物中分离出近70个木脂素成分,并着重介绍木脂素成分降低谷丙转氨酶的研究结果以及其他主要活性。并对国内五味子科植物的分布和资源利用作了介绍.     

Comparative morphology of the leaf epidermis in Schisandra (Schisandraceae)

The leaf epidermis of 127 samples representing ten species within Schisandra Michaux. and one species of the related genus, Kadsura Kaempf. ex Juss., was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandra , such as pattern of epidermal cells, type of stomata, shape of guard cell pairs and cuticular ornamentation, are usually constant within species and thus make good characters for studying the relationship between and within genera. A new character, rim number, of the outer stomatal rim in the genus is introduced. It is shown that double outer stomatal rims occur only in evergreen Schisandra species, whereas a single rim occurs in deciduous species. This character supports the classification of Schisandra into two subgenera based on habit and androecial organs. This classification is also supported by additional morphological and molecular taxonomic characters. Kadsura coccinea (Lem.) Smith A. C. is the most primitive taxon in the related genus Kadsura . The outer stomatal rim of this species also has double rims. Combined with morphological and molecular evidence, this suggests that Schisandra and Kadsura are closely related and may share a recent common ancestor.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 39–56.     

Morphological characters of leaf epidermis in schisandraceae and their systematic significance

The leaf epidermis of 23 species belonging to 2 genera within Schisandraceae was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandraceae, such as shape of epidermal cells, type of stomata, and cuticular ornamentation, are usually constant within species and thus helpful for elucidating the relationship between and within genera. Leaf epidermal cells are usually irregular or polygonal in shape. The patterns of anticlinal walls are straight, sinuolate, sinuous or sinuate. The stomatal apparatus belong to paracytic or laterocytic type and the latter is subdivided into various subtypes based on the number and arrangement of subsidiary cells. Under scanning electron microscopy observation, the cuticular membrane is often striated, sometimes squamulate or granular; the inner margin of the outer stomatal rim is nearly smooth or denticulate. Evidences from shape of epidermal cells, patterns of cuticular intrusions between the ends of each guard cell of a pair and distribution of stomatal apparatuses support the viewpoint thatKadsura is more primitive thanSchisandra. Study on leaf epidermis also shows thatKadsura interior deserves the rank of a distinct species and the treatment of the evergreen groups, includingS. propinqua andS. plena, as distinct from the deciduous species of the genus is quite natural.     

五味子科植物花形态及其系统学意义

用光镜和扫描电镜观察了五味子科Ｓｃｈｉｓａｎｄｒａｃｅａｅ２３种、６变种和１变型的９１份花粉的形态。其中５ 、４变种和１变型的花粉形态为首次报道。该科花粉为单花粉,异极,辐射对称,扁球形至超扁球形,其萌发沟的排列格局在被子植物中是独一无二的。若按萌发沟的数目划分,该科花粉可分为３沟型和６沟型两个类型。外壁为网状纹饰,网眼的大小和网脊的宽度与花的形态有一定相关性。研究结果不支持Ｐｒａｇｌｏｗｓｋｉ对     

Pollen morphology of the Schisandraceae and its systematic significance

Pollen morphology of 91 samples, representing 23 species, 6 varieties and 1 form of the Schisandraceae, was examined under light microscope (LM) and scanning electron microscope (SEM). Palynological data of 5 species, 4 varieties and 1 form are reported for the first time. The pollen grains are monads, heteropolar, radiosymmetric and oblate to peroblate in shape. Arrangement of their colpi is unique in angiosperms. The pollen grains can be divided into two types according to the number of their clopi: the tricolpate pollen type and the hexacolpate pollen type. The exine sculpturing is reticulate. The size of luminae and the breadth of muri are correlated with the floral morphology of the hisandraceae to a certain extent. Our results do not support the division of the pollen grains of the Schisandraceae into 4 types according to the size of inae as proposed by Praglowski. There is also still room of discussion about his conclusion that the pole where the three longer colpi converge is the distal pole. The evolutionary trends of the pollen grains of the Schisandraceae seem to be from fewer to more in the number of colpi, and from smaller to larger in the size of luminae. The view that Kadsura is more primitive than Schisnadra is not supported by palynological data; on the contrary, Kadsura seems to be more advanced than Schisandra at the general level of evolution. The two genera may be closely related, and might have originated from a common ancestor and then evolved parallelly along two different routes.     

基于ITS与 trn L-F序列探讨槭树科的系统发育

报道了槭树科41种(其中槭属39种)植物的 trn L-F和ITS序列(其中部分种的ITS序列为重新测定),以期通过分子手段对槭树科内部尤其是复杂的槭属的系统发育关系进行重建.以无患子科和七叶树科为外类群,基于对57个种单独的ITS序列(包括从GenBank下载的16种的序列)、41种 trn L-F序列及41种两者序列的联合数据,分别采用最大简约法(Maximum Parsimony Method)和邻接法(Neighbor-Joining Method)对槭树科的系统发育进行了分析.结果显示,整个槭树科为一单系类群;金钱槭位于槭树科的基部;但由于云南金钱槭( Dipteronia dyerana )聚在了槭属内部,认为金钱槭属和槭属均可能是非单系类群;槭属内组间关系的支持率普遍较低,但多数组的组内关系得到了较好的支持.将两个片段结合比单独的ITS或 trn L-F分析能更好地解决槭属内部的系统关系,其中sect.Palmata 和sect.Microcarpa ,sect.Platanoidea 、sect.Lithocarpa 和sect.Macrophylla ,sect.Integrifolia 、sect.Trifoliata 和sect.Pentaphylla ,以及sect.Acer 、sect.Goniocarpa 和sect.Saccharina (sensu Ogata)的组间亲缘关系得到了一定的支持,但对其中部分组的划分可能应做进一步调整.重新评价了徐廷志系统中对sect.Rubra 和sect.Saccharodendron 的处理.     

基于ITS与trnL—F序列探讨槭树科的系统发育

报道了槭树科41种（其中槭属39种0植物的trnL-F和ITS序列（其中部分种的ITS序列为重新测定），以期通过分子手段对槭树科内部尤其是复杂的槭属的系统发育关系进行重建。以无患子科和七叶树科为外类群，基于对57个种单独的ITS序列（包括从GenBank下载的16种的序列），41种trnL-F序列及41种两序列的联合数据，分析采用最大简约法（Maximum Parsimony Method)和邻接法（Neighbor-Joining Method)对槭树科的系统发育进行了分析。结果显示，整个槭树科为一单系类群；金钱槭位于槭树科的基部；但由于云南金钱槭（Dipteronia dyerana)聚在槭属内部，认为金钱槭属和槭属均可能是非单系类群；槭属内组间关系的支持率普遍较低，但多数组的组内关系得到了较好的支持。将两个片段结合比单独的ITS或trnL-F分析能更好地解决槭属内部的系统关系，其中sect,Palmata和sect.Micrcarpa,sect,Platanoidea,sect,Lithocarpa和sect.Macrophylla,sect,Integrifolia.Trifoliata和sect Pentaphylla,以及sect.Acer,sect.Goniocarpa和sect.Saccharina(sensu Ogata）的组间亲缘关系得到了一定的支持，但对其中部分组的划分可能应做进一步调整。重新评价了徐廷志系统中对sect.Rubra和sect.Saccharodendron的处理。     

南五味子属花的形态及其系统学意义

借助扫描电镜观察了南五味子属(Kadsura)中黑老虎(K.coccinea)、南五味子(K.japonica)和狭叶南五味子(K.angustifolia)雄花和雌花的形态发生过程, 三种植物花发育早期相似而发育后期出现分异。根据花形态发生早期的相似性, 支持南五味子属为单系起源;根据花形态发生后期的分异状态, 支持将南五味子属分为离蕊南五味子亚属(Subgenus Cosbaea)和南五味子亚属(Subgenus Kadsura)。借助扫描电镜观察了短梗南五味子(K.borneensis)和披针叶南五味子(K.lanceolata)花的形态, 将南五味子和狭叶南五味子的花与短梗南五味子和披针叶南五味子的花比较, 赞成在南五味子亚属下设南五味子组(Section Kadsura)和南洋南五味子组(Section Sarcocarpon)。根据在狭叶南五味子中观察到的两性花痕迹和在五味子属(Schisandra)东亚五味子(S.elongata)花中观察到的两性花, 本文认为现存五味子科(Schisandraceae )植物的单性花可能由具两性花结构的祖先演化而来。根据五味子科植物柱头与八角科(Illiciaceae)植物柱头的相似性, 不支持将五味子科从八角目(Illiciales)中分出而成立五味子目(Schisandrales)的观点。     

五味子属（五味子科）分类系统的初步修订

通过对前人提出的五味子属（五味子科）分类系统的点评,根据五味子属植物的33个形态性状及其性状分析,经过分支分析,提出一个新的五味子属分类系统：将五味子属分为五味子亚属和团蕊五味子亚属;将五味子亚属分为多蕊五味子组、少蕊五味子组、中华五味子组和五味子组。编排了分亚属、组和种的检索表,对每个种上分类等级均列出其模式、异名和代表种。     

《秦岭植物志》种子植物补遗--被子植物门的双子叶植物(1)

报道了《秦岭植物志》遗漏的26种(含3个种下分类群)植物,26种植物分属于17属,13科。其中大血藤科(Sargentodoxaceae)和铁青树科(Olacaceae)为遗漏科,大血藤科的大血藤属(Sargentodoxa)、铁青树科的青皮木属(Schoepfia)、五味子科的南五味子属(Kadsura)、樟科的黄肉楠属(Actinodaphne)和虎耳草科的钻地风属(Schizophragma)为遗漏属。     

Phylogeny of Schisandraceae based on morphological data: evidence from modern plants and the fossil record

Schisandraceae are traditionally subdivided in two genera, Schisandra and Kadsura, based on differences in the organisation of the floral receptacle, the carpels, and the presence or absence of a ``pseudostigma'. Recently, phylogenetic analyses utilizing ITS sequence data and morphological data resulted in incongruent tree topologies, with the morphological trees suggesting monophyly of the two genera, whereas ITS trees did not resolve Schisandra and Kadsura as monophyletic clades. In the present paper we study seed morphology and leaf epidermal features of 22 species of Schisandraceae in order to provide additional data for a morphological data matrix. Seed morphological characters are highly homoplastic and do not yield further evidence for monophyly of the two genera. Instead, a number of characters appear to support sister group relationships between taxa within the genera, such as, for instance, for K. coccinea and K. scandens, both of which have large seeds along with a multi-layered mesotesta. Considering leaf epidermal characteristics, species of Kadsura were found to be consistently amphistomatic, whereas species of Schisandra are always hypostomatic. Phylogenetic analysis using the extended data matrix resulted in weakly supported Kadsura and Schisandra clades with five and four synapomorphies indicating monophyly of Kadsura and Schisandra, respectively. Fossils ascribed to Schisandraceae date back to the Late Cretaceous. These are tri-and hexacolpate pollen types displaying a combination of features found in modern Schisandraceae and partly also in Illiciaceae. Leaf remains from this period are poorly preserved and difficult to ascribe to Schisandraceae because of the lack of synapomorphies for the family. In the Early Cainozoic, leaf and seed remains from North America and Europe unambiguously belong to the family. Seeds from the Eocene of North America show some similarities to the modern Schisandra glabra from North America, while fossils from Europe show more similarities to modern Asian species.     

Systematics of the cyclostome subfamilies of braconid parasitic wasps (Hymenoptera: Ichneumonoidea): a simultaneous molecular and morphological Bayesian approach

Phylogenetic relationships among 95 genera collectively representing 17 of the 18 currently recognized cyclostome braconid wasp subfamilies were investigated based on DNA sequence fragments of the mitochondrial COI and the nuclear 28S rDNA genes, in addition to morphological data. The treatment of sequence length variation of the 28S partition was explored by either excluding ambiguously aligned regions and indel information (28SN) or recoding them (28SA) using the 'fragment-level' alignment method with a modified coding approach. Bayesian MCMC analyses were performed for the separate and combined data sets and a maximum parsimony analysis was also carried out for the simultaneous molecular and morphological data sets. There was a significant incongruence between the two genes and between 28S and morphology, but not for morphology and COI. Different analyses with the 28SA data matrix resulted in topologies that were generally similar to the ones from the 28SN matrix; however, the former topologies recovered a higher number of significantly supported clades and had a higher mean Bayesian posterior probability, thus supporting the inclusion of information from ambiguously aligned regions and indel events in phylogenetic analyses where possible. Based on the significantly supported clades obtained from the simultaneous molecular and morphological analyses, we propose that a total of 17 subfamilies should be recognized within the cyclostome group. The subfamilial placements of several problematic cyclostome genera were also established.     

Molecular identification of Schisandra chinensis and its allied species using multiplex PCR based on SNPs

Schisandra chinensis (Turcz.) Baill. is a climbing woody vine found over a wide geographical range in northeast Asia. Its red five-flavored fruits have been used as drugs and food. In China and Korea, the dried fruit of S. chinensis has been used for a long time as a traditional medicine. Three species of the family Schisandraceae, S. chinensis, S. repanda, and Kadsura japonica, are distributed mainly in southern Korea. While S. chinensis is broadly distributed, S. repanda and K. japonica are restricted to the southern islands of Korea, especially Jeju. The amplification refractory mutation system (ARMS), one of molecular genotyping technique, is a simple and rapid method for detecting point mutations, restriction fragment length polymorphisms, and small deletions or insertions in a DNA sequence. This PCR-based technique utilizes a primer designed to react only when it anneals to specific target sequences in a DNA sample. In the present study, we analyzed rbcL of the chloroplast gene and ITS from S. chinensis and two related taxa, S. repanda and K. japonica, and developed a specific marker for distinguishing S. chinensis from the related taxa based on SNPs using multiplex PCR. Result showed that two species-specific fragments with 230 bp of rbcL and 278 bp of ITS region were additionally amplified only in all of individuals of S. chinenesis using multiplex PCR method. Here, we describe the details of the protocol for the molecular identification of S. chinensis and other related species. Our results cover five populations of S. chinensis in Korea.