睡莲科的核型分析及其分类学位置的探讨

本文对睡莲科6属6种代表植物的核型进行了研究,并探讨了它的分类学位置。结果如下：莲2n＝16=9sm+4m+3st;王莲2n＝24=8sm+8m+8T,蓝睡莲2n＝28,可配成14对,染色体小,第l号染色体上有2条随体;萍蓬草2n＝34＝18m+16sm;芡实2n=58,可配成29对,染色体小,第l号染色体有2条随体,莼菜2n＝72,可配成36对,染色体按大小可分成大,中、小三个类别。除莲外,其它5种植物的核型为首次报道。莼菜的体细胞染色体数目2n＝72和国外报道的2n＝80不相一致。莲的染色体以及形态学特征和其它睡莲科分类群显著不同,可将其从睡莲科中独立出来,并成立莲科和莲目。原归属于睡莲科的分类群仍组成睡莲目,并分别置于莼菜科和睡莲科。     

Phylogeny of the Nymphaeaceae inferred from

The Nymphaeaceae are one of the most important taxa for understanding the origin and evolution of primitive angiosperms due to its basal position in the cladograms of the angiosperms recently given based both on several gene sequences and on morphological characters, but phylogenetic relationships within the family have not as yet been resolved. The sequences of nrDNA ITS region of 11 species representing seven genera of the Nymphaeaceae and one outgroup, Ceratophyllum demersum, were used to reconstruct the phylogeny of the family using PAUP4.0b4A. Three most parsimonious trees (Length=1125, CI=0.7618 and RI=0.7214) were obtained. In the consensus tree: (1) Nelumbo was basal to the other genera with a bootstrap value of 100% and occupied an isolated position, so it could be separated from the Nymphaeaceae and placed in its own family, Nelumbonaceae, and its own order, Nelumbonales; (2) Nuphar was monophyletic, nested in the basal position of clade II and was strongly supported to be the sister group of the other genera (excluding Nelumbo) of the Nymphaeaceae, suggesting that its traditional inclusion in the Nymphaeaceae should be maintained; (3) Cabomba and Brasenia were sister group and formed a subclade with a bootstrap value of 99%, indicating their close affinity; (4) Nymphaea was strongly supported to be the sister group of a subclade comprising Euryale and Victoria with a bootstrap value of 94%, indicating that they should be placed in the Nymphaeaceae.     

Vessels in the roots of Barclaya rotundifolia (Nymphaeaceae)

Scanning electron microscope (SEM) studies of paraffin-sectioned material of stems and roots of Barclaya rotundifolia Hotta revealed perforations on tracheary elements of roots, but not on those of stems. End walls of vessels are identical with lateral walls except for the presence of perforations. Perforations can only be clearly revealed with SEM, and this method is advocated for further study of tracheary elements of Nymphaeaceae in particular, and primary xylem of ∗∗∗angiosperms in general. Vessel presence may be related to the habitat of this species, which unlike other members of the Nymphaeaceae (sensu stricto) has only aerial leaves and a rhizomatous system that is not inundated for prolonged periods.     

睡莲科的系统发育:核糖体DNA ITS区序列证据

以金鱼藻Ceratophyllum demersum为外类群,使用PAUP4.0b4A软件对睡莲科Nymphaeaceae植物7属11个代表种的ITS区序列进行了系统发育分析。采用最大简约法分析获得了3个最简约树,步长为1125,一致性指数(CI)和维持性指数(RI)值分别为0.7618和0.7214。利用3个最简约树获取严格一致树。结果表明:(1)莲属Nelumbo位于系统树的基部,自展支持率为100%,可从睡莲科中独立出来成立莲科Nelumbonaceae和莲目Nelumbonales;(2)萍蓬草属Nuphar是一单系类群,位于分支Ⅱ的基部,并和睡莲科其他属(不包括莲属)植物聚在一起构成姐妹群,故萍蓬草属仍应置于睡莲科中;(3)水盾草属Cabomba和莼菜属Brasenia聚成一小支并构成姐妹群,自展支持率为99%,说明这两属之间亲缘关系较近;(4)睡莲属和芡实属Euryale、王莲属Victoria聚成一小支并构成姐妹群,自展支持率为94%,说明三者亲缘关系较近,仍应置于睡莲科中。     

睡莲科三属植物叶的比较解剖学研究

报道了睡莲科的睡莲属、莼菜属和萍蓬草属三属植物叶的比较解剖学特征。结果表明,莼菜属叶柄仅具一个维管束,其余二个属均有数个维管束,且维管束为有限的、星散排列,这一特征与单子叶植物的很相似;叶柄基本组织中,除萍蓬草属无气道外,其它两个属均有气道;在睡莲属和萍蓬草属中存在星状石细胞,而莼菜属中则无此细胞;莼菜属和萍蓬草属具腺毛,睡莲属无腺毛。由于睡莲科此三属植物叶的解剖学特点存在很大的差异,因此认为睡莲科的这三属可能来源于不同的祖先,因此建议将莼菜属和萍蓬草属从睡莲科中分离出来。分别独立为莼菜科和萍蓬草科。     

Perianth organs in Nymphaeaceae: comparative study on epidermal and structural characters

Journal of Plant Research - The perianth organs of six species of Nymphaeaceae, representing Euryale, Nymphaea and Victoria, were studied on the basis of macroscopical, micromorphological, and...     

Phylogenetic analysis of Nymphaeales using fast-evolving and noncoding chloroplast markers

The Nymphaeales (water-lilies and relatives) represent one of the earliest branching lineages of angiosperms and comprise about 70 aquatic species. Here, we present a comprehensive study of phylogenetic relationships within the Nymphaeales from a dataset containing 24 representatives of the order, including all currently recognized genera and all subgenera of the genus Nymphaea , plus 5 outgroup taxa. Nine different regions of the chloroplast genome − comprising spacers, group II introns, a group I intron, and a protein coding gene − were analysed. This resulted in a character matrix of 6597 positions and an additional 369 characters obtained from coded length mutations. Maximum parsimony and Bayesian analyses of the complete dataset yielded congruent, fully resolved and well-supported trees. Our data confirm the monophyly of the Cabombaceae but do not provide convincing support for the monophyly of Nymphaeaceae with respect to Nuphar . Moreover, the genus Nymphaea is inferred to be paraphyletic with respect to Ondinea , Victoria and Euryale . In fact, the Australian endemic Ondinea forms a highly supported clade with members of the Australian Nymphaea subgenus Anecphya . In addition, Victoria and Euryale are inferred to be closely related to a clade comprising all night-blooming water-lilies ( Nymphaea subgenera Hydrocallis and Lotos ). An experimental approach showed taxon sampling to be of influence on the nodes reconstructed in core Nymphaeaceae. The results underscore that more diverse genera, if not clearly known to be monophyletic, should be represented by all major lineages.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 141–163.     

越南杜鹃花科新记录——深裂树萝卜(英文)

报道了越南杜鹃花科(Ericaceae)1新记录种——深裂树萝卜(Agapetes lobbii C.B.Clarke)。该种产自越南北部莱州省(Lai Chau)的Sin Ho地区,凭证标本保存在越南科学技术院生态和生物资源研究所标本馆(HN)。     

中国睡莲科植物种子胚及其营养组织的研究

研究了中国睡莲科的3属3种即芡（Euryale ferox Salisb.）、睡莲（Nymphaea tetragona Georgi）和萍蓬草（Nuphar pumilum (Hoffm.) DC.）的胚、胚乳及外胚乳。它们的胚的共同特点是苗端发育良好,特别是胚芽具2～4枚胚芽叶,而根端滞育。三个种皆既有胚乳,又有外胚乳,但在成熟种子中作为胚的营养组织则胚乳弱育而外胚乳发达。最后对上述特点在系统发生中的意义进行了讨论。     

Comparative Leaf Anatomy of Nymphaeaceae (s. l.)

Leaf anatomy of six genera was investigated. The petiole of Brasenia schreberi possesses a pair of bundles, the other five possess closed, scattered vascular bundles, resembling those of the Monocotyledons. True vessels are absent. Air passages are present in the ground tissue of the petiole except for Nuphar pumilum. Leaves are all dorsiventral. Stomatal apparatuses of the Ranunculaceous type are generally confined to the upper surface of the lamina, but fewer stomata are found on the lower surface of the lamina of Nelumbo nucifera. The stomatal apparatuses of Nelumbo nucifera are of the Haplocheilic type in development, but of the Ranunculaceous type at maturity. Hydathodes and sclerenchymatous idioblasts are present only in Nelumbo nucifera. Glandular hairs and hydropotens occur on the lower surface of the lamina in all the investigated species except Nelumbo nucifera. Astrosclereids are present in all the taxa except Brasenia schreberi and Nelumbo nucifera. According to observations we consider that the genus Brasenia is a primitive one in Nymphaeaceae and it has a close phylogenetic relationship with Cabomba. They may be treated as an independent family, Cabombaceae, which is better included in the order Nymphaeales. Nymphaea, Nuphar, Euryale and Victoria are closely related to one another and should be retained in the family Nymphaeaceae. Nelumbo shows, specialized features distinct from those of the rest of the Nymphaeaceae. It should be segregatednot only as a distinct family but also as a distinct order, Nelumboales.     

Pollen ontogeny in Brasenia (Cabombaceae, Nymphaeales)

Brasenia is a monotypic genus sporadically distributed throughout the Americas, Asia, Australia, and Africa. It is one of eight genera that comprise the two families of Nymphaeales, or water lilies: Cabombaceae (Brasenia, Cabomba) and Nymphaeaceae (Victoria, Euryale, Nymphaea, Ondinea, Barclaya, Nuphar). Evidence from a range of studies indicates that Nymphaeales are among the most primitive angiosperms. Despite their phylogenetic utility, pollen developmental characters are not well known in Brasenia. This paper is the first to describe the complete pollen developmental sequence in Brasenia schreberi. Anthers at the microspore mother cell, tetrad, free microspore, and mature pollen grain stages were studied using combined scanning electron, transmission electron, and light microscopy. Both tetragonal and decussate tetrads have been identified in Brasenia, indicating successive microsporogenesis. The exine is tectate-columellate. The tetrad stage proceeds rapidly, and the infratectal columellae are the first exine elements to form. Development of the tectum and the foot layer is initiated later during the tetrad stage, with the tectum forming discontinuously. The endexine lamellae form during the free microspore stage, and their development varies in the apertural and non-apertural regions of the pollen wall. Degradation of the secretory tapetum also occurs during the free microspore stage. Unlike other water lilies, Brasenia is wind-pollinated, and several pollen characters appear to be correlated with this pollination syndrome. The adaptive significance of these characters, in contrast to those of the fly-pollinated genus Cabomba, has been considered. Brasenia does not produce pollenkitt nor develop tectal microchannels as does Cabomba. Instead, the discontinuity of the tectum reduces the amount of sporopollenin in the wall, which may allow for more effective wind dispersal. The importance of reassessing palynological characters in light of new ontogenetic data and the phylogenetic implications of this reevaluation are also discussed.     

Seedlings of Nymphaeaceae

Seedlings of Nymphaea coerulea, N. lotus and Nuphar luteum were examined for monocoty-ledonous and dicotyledonous characters. The cotyledonary formation could be interpreted either as a single bilobed structure or as two separate cotyledons. However, the presence of an operculum and the recognition of a coleoptile and mesocotyl similar to those of grasses and sedges gave grounds for including Nymphaeaceae in the monocotyledons. The form of the hypocotyl and radicle and the leaf-type supported a position, as a specialized family, in or close to the Helobiae.     

Recent and Tertiary Seeds of the Nymphaeaceae sensu lato with a Revision of Brasenia ovula (Brong.) Reid and Chandler

The nine extant genera within the Nymphaeaceae s l (Water Lilies)are shown to be separable from one another on the basis of seedmorphology In particular, epidermal detail, vertical sectionof the testa and relative position of the micropyle and hilumare diagnostic for each genus These features are consideredin connection with current systematic treatments of this familyFossil seeds are reviewed at the generic level and many of theseeds previously assigned to Brasenia ovula (Brong ) Reid andChandler are shown to belong to an extinct genus of Nymphaeaceaeintermediate between the currently accepted families Cabombacaceaeand Nymphaeaceae s s They are redescribed as Sabrenia chandleraegen et sp nov. The British Tertiary fossils Brasenia spinosaChandler, Palaeonymphaea eocenica Chandler emend and ?Nymphaealiminis sp nov are described Fossil material of Carpolithesovulum Brongniart 1822a, Brasenia victoria (Casp ) Weberbauer1893, B teumeri Kirchheimer 1935, B tenuicostata Nikitin 1965and Nymphaea arethusae Grambast 1962 has been studied for comparativepurposes The use of ‘?’ before the extant genericname is advocated when all features of a fossil indicate thatit may be placed in a living genus but when certain additional,critical features of the living genus are lacking on the fossil. Nymphaeaceae, water lilies, seeds, palaeobotany, Tertiary, Sabrenia chandlerae gen et sp nov Nymphaea liminis sp nov, Brasenia ovula     

Ultraviolet reflectance and absorption patterns in flowers of Nymphaea alba L., Nymphaea candida presl and Nuphar lutea (L.) Sm. (Nymphaeaceae)

Ultraviolet (u.v.) patterns of flowers of Nymphaeaceae as an important colour component for flower-visiting insects have been totally neglected in studies of their floral biology. The three day-flowering species studied show distinct u.v. patterns. Both Nymphaea species show hogh absorption in the centre of the flowers and u.v.-reflective thecae. In Nymphaea alba L. a radial u.v. stripe reflection pattern is visible on the stigmatic disc in u.v. photographs, which is weakly developed in Nymphaea candida Presl flowers. The flowers of Nuphar show the most distinct u.v. reflection pattern in the form of a circle formed by the u.v.-reflective tips and filaments of the stamens, in contrast to the shiny petals and u.v.-absorptive stigmatic disc, thecae and sepals.     

The vulnerability of habitats to plant invasion: disentangling the roles of propagule pressure,time and sampling effort

Aim To quantify the vulnerability of habitats to invasion by alien plants having accounted for the effects of propagule pressure, time and sampling effort. Location New Zealand. Methods We used spatial, temporal and habitat information taken from 9297 herbarium records of 301 alien plant species to examine the vulnerability of 11 terrestrial habitats to plant invasions. A null model that randomized species records across habitats was used to account for variation in sampling effort and to derive a relative measure of invasion based either on all records for a species or only its first record. The relative level of invasion was related to the average distance of each habitat from the nearest conurbation, which was used as a proxy for propagule pressure. The habitat in which a species was first recorded was compared to the habitats encountered for all records of that species to determine whether the initial habitat could predict subsequent habitat occupancy. Results Variation in sampling effort in space and time significantly masked the underlying vulnerability of habitats to plant invasions. Distance from the nearest conurbation had little effect on the relative level of invasion in each habitat, but the number of first records of each species significantly declined with increasing distance. While Urban, Streamside and Coastal habitats were over‐represented as sites of initial invasion, there was no evidence of major invasion hotspots from which alien plants might subsequently spread. Rather, the data suggest that certain habitats (especially Roadsides) readily accumulate alien plants from other habitats. Main conclusions Herbarium records combined with a suitable null model provide a powerful tool for assessing the relative vulnerability of habitats to plant invasion. The first records of alien plants tend to be found near conurbations, but this pattern disappears with subsequent spread. Regardless of the habitat where a species was first recorded, ultimately most alien plants spread to Roadside and Sparse habitats. This information suggests that such habitats may be useful targets for weed surveillance and monitoring.     

Genetically based polymorphisms in morphology and life history associated with putative host races of the water lily leaf beetle,Galerucella nymphaeae

A host race is a population that is partially reproductively isolated from other conspecific populations as a direct consequence of adaptation to a specific host. The initial step in host race formation is the establishment of genetically based polymorphisms in, for example, morphology, preference, or performance. In this study we investigated whether polymorphisms observed in Galerucella nymphaeae have a genetic component. Galerucella nymphaeae, the water lily leaf beetle, is a herbivore which feeds and oviposits on the plant hosts Nuphar lutea and Nymphaea alba (both Nymphaeaceae) and Rumex hydrolapathum and Polygonum amphibium (both Polygonaceae). A full reciprocal crossing scheme (16 crosses, each replicated 10 times) and subsequent transplantation of 1,001 egg clutches revealed a genetic basis for differences in body length and mandibular width. The heritability value of these traits, based on midparent-offspring regression, ranged between 0.53 and 0.83 for the different diets. Offspring from Nymphaeaceae parents were on average 12% larger and had on average 18% larger mandibles than offspring from Polygonaceae parents. Furthermore, highly significant correlations were found between feeding preference of the offspring and the feeding preference of their parents. Finally, two fitness components were measured: development time and survival. Development time was influenced by diet, survival both by cross type and diet, the latter of which suggest adaptation of the beetles. This suggestion is strengthened by a highly significant cross x diet interaction effect for development time as well as for survival, which is generally believed to indicate local adaptation. Although no absolute genetic incompatibility among putative host races was observed, survival of the between-host family offspring, on each diet separately, was lower than the survival of the within-host family offspring on that particular host. Survival of offspring of two Nymphaeaceae parents was about two times higher on Nymphaeaceae than on Polygonaceae, whereas survival of offspring of two Polygonaceae parents was 11 times higher on Polygonaceae than on Nymphaeaceae (based on untransformed data). Based on these results, we conclude that genetically determined polymorphisms in morphology and feeding preference exist in G. nymphaeae, resulting in differential performance. Furthermore, in each diet separately, offspring of between-host family crosses were less fit than offspring of within-host family crosses. These results support the hypothesis that within this species two host races can be distinguished.     

Megagametophyte development of Nymphaea nouchali Burm. f. (Nymphaeaceae)

The development of reproductive cells has been analysed in Nymphaea nouchali , family Nymphaeaceae, subclass Magnoliidae. Our observations on embryo sac development differ from those in the literature. Megasporogenesis results in a triad of one micropylar cell and two innermost megaspores. Embryo sacs derive from a single functional megaspore and are tetranucleate. Megasporogenesis is monosporic and of the Polygonum type, but mega-gametogenesis is of Oenothera type.     

Pollination of Nymphaea lotus (Nymphaeaceae) by Rhinoceros Beetles and Bees in the Northeastern Ivory Coast

Abstract: Detailed observations of the pollination biology of Nymphaea lotus were conducted by studying populations growing in seasonal ponds in the Comoé National Park (NE Ivory Coast). Flowers of N. lotus are protogynous and anthesis lasts for 4 to 5 days. Flowers open at sunset and close in the morning hours, thereby revealing a high variability in timing. For the first time, heat production in flowers could be observed in the genus Nymphaea. Flowers of N. lotus were visited by Ruteloryctes morio, an endemic dynastid beetle during the night. Beetles copulated in the flowers and fed on flower parts, but were less effective pollinators than several bee species that visited flowers in the early morning. Nymphaea lotus thus seems to be adapted to pollination by both nocturnal beetles and diurnal bees.     

Do tracheid microstructure and the presence of minute crystals link Nymphaeaceae,Cabombaceae and Hydatellaceae?

Original scanning electron microscopy (SEM) observations are presented for stems of Brasenia schreberi and Cabomba caroliniana of Cabombaceae and three species of Trithuria of Hydatellaceae. End walls of stem tracheids of Brasenia have the same peculiar microstructure that we have reported in Barclaya, Euryale, Nuphar, Nymphaea (including Ondinea) and Victoria of Nymphaeaceae. This feature unites Cabombaceae with Nymphaeaceae. The minute rhomboidal crystals on the surfaces of stellate parenchyma cells of Brasenia reported by Solereder (1906. Oxford: University Press), but not noticed since, are figured. They are like the minute crystals of the often‐mentioned astrosclereids of Nymphaeaceae. Neither of these two features has been observed in Hydatellaceae. If the absence of these two features can be confirmed, the reason may be more related to ecology, development, habit and anatomical organization than to degree of phylogenetic relationship as shown by molecular studies. Anatomical observations on the stem anatomy of Trithuria are offered on the basis of paraffin sections prepared for a paper by Cheadle & Kosakai (1975. American Journal of Botany 62: 1017–1026); that study is notable for a discrepancy between an illustration of a specialized vessel element on the one hand and tabular data indicating long scalariform perforation plates on the other. Long scalariform perforation plates are mostly found in scalariformly pitted vessels of monocots, whereas the tracheary elements of Trithuria mostly have helical or annular thickenings. We were unable to demonstrate the presence of vessels in Hydatellaceae. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 572–582.     

Endoplasmic reticulum derived decorated tubules in the sieve elements ofNymphaea

Summary Bundles of decorated tubules found in the sieve elements ofNymphaea have been studied with the transmission electron microscope. Comparatively straight tubules (100 nm in diameter) arise from the endoplasmic reticulum during early stages of sieveelement development and subsequently associate into bundles of up to 100 tubules that parallel the longitudinal cell axis. From the start of their formation the tubules are structurally distinct from other ER profiles due to their dense decoration with particles. High magnifications reveal an orderly array of the particles (about 24 surround a 100 nm tubule) and suggest a modification of their membrane so that it is no longer dissolvable into a regular three-layered structure. Later during sieve-element ontogeny the decorated tubules get invaginated by smooth ER membranes, thereby squeezing out the intratubular (extracytoplasmic) space. As a result a double mantle is formed that surrounds a plasmatic cylinder. Decorated 100 nm tubules with inner membranes are present in enucleate mature sieve elements ofNymphaea alba andN. tuberosa. Considerably larger tubules (about 200 nm in diameter) were found inN. Candida andN. tetragona and occasionally also inNuphar and Barclaya, two other genera from the same family. The decoration of the tubules and their subsequent invagination by smooth membranes are discussed with respect to the controlled autolysis of sieve elements.Dedicated to Prof. Dr. Dr. h.c. Eberhard Schnepf on the occasion of his retirement