Phylogenetic relationships in Nuphar (Nymphaeaceae): evidence from morphology,chloroplast DNA,and nuclear ribosomal DNA

The genus Nuphar consists of yellow-flowered waterlilies and is widely distributed in north-temperate bodies of water. Despite regular taxonomic evaluation of these plants, no explicit phylogenetic hypotheses have been proposed for the genus. We investigated phylogenetic relationships in Nuphar using morphology and sequences of the chloroplast gene matK and of the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA. Two major lineages within Nuphar are consistently resolved with the morphological and molecular data sets. One lineage comprises New World taxa and the other represents a primarily Old World lineage. Relationships within the major lineages were poorly resolved by morphology and ITS, yet certain relationships were elucidated by all analyses. Most notable is the strong support for a monophyletic lineage of dwarf taxa and the alliance of the North American N. microphylla with the Eurasian taxa. Minor discordance between the independent cladograms is accounted for by hybridization. The common taxonomic practice of uniting all North American and Eurasian taxa under one species is not supported phylogenetically.     

Analysis of the fruit surfaces in Bupleurum L. (Umbelliferae) with SEM

20 Bupleurum L. taxa were investigated for micromorphological typology of the fruit surfaces with SEM. Considerably different types of sculpturing at species and subspecies level and similarities between related taxa were observed. A dichotomic key was prepared using the diagnostic surface patterns. Reticulate (sect. Bupleurum), rugose (sect. Aristata Godron subsect. Aristata) and rugulose-colliculate-verrucate (subsect. Juncea Briq.) types of surface structures were distinguished. Micromorphological characteristics may be valuable additional taxonomical criteria for the delimitation of infrageneric groups of species.     

Systematic analysis of Dendrobium Swartz section Dendrocoryne in the Australian region

Dendrobium sect. Dendrocoryne is a difficult taxonomic group of allied species and species complexes, occurring on the eastern Australian coastline, Lord Howe Island and New Caledonia. Significant morphological characters were assessed from all key areas of the distribution range. Patterns of variation between species, varieties and hybrids were investigated by principal coordinate analysis. The analyses affirm 12 species in the sect. Dendrocoryne described by Schlechter (1912, 1982) and Dockrill (1969, 1992), including a cluster of small statured D. gracilicaule, D. adae, D. fleckeri, and D. finniganense, and three variable species complexes – D. speciosum, D. kingianum and D. tetragonum. Recently described D. finniganense and D. callitrophilum are interpreted as members of the section. Phylogenetic relationships are presented based on parsimony analysis of 31 morphological characters. Analyses do not support proposals to recognize new genera for D. tetragonum, D. callitrophilum and D. aemulum, or to create new species within D. tetragonum, D. jonesii, D. speciosum and D. kingianum. The phylogeny indicates that rainforest taxa are earlier lineages and xerophytic taxa are more derived.     

Patterns of allozyme variation in Nordic Pilosella

Patterns of allozyme variation have been investigated in 101 clones representing most of the morphological variation (and taxa) found in the amphi-apomictic genus Pilosella Hill. in the Nordic countries. Levels of variation were very high, both within narrowly defined taxa and within the whole data-set, but the correlation between a morphology-based taxonomy and patterns of allozyme variation was very weak. Almost all enzyme bands were found in more than one taxon and the differences between broadly defined species in terms of the frequency of bands were generally small. Cluster analyses did not perfectly recover any taxon but there were some tendencies for samples of some broadly defined taxa (e.g. P. officinarum s. l., and P. cymosa ssp. cymosa) to cluster together. It is concluded that there is considerable gene-flow among virtually all Nordic morpho-types of Pilosella and this is considered an argument for applying a very broad species concept in this genus.Financial support for the present study has been recieved from Stiftelsen Harald E. Johanssons minnesfond at the Royal Swedish Academy of Sciences.     

Nutlet morphology in Hemigenia R.Br. and Microcorys R.Br. (Lamiaceae)

Nutlets of Hemigenia R.Br. and Microcorys R.Br. were examined using SEM. Significant variation, mainly useful at the infrageneric level, was found in nutlet shape, nature of the attachment scar, nature of surface sculpturing, exocarp cell shape and sculpturing, and nature of the indumentum. Typical nutlets are ovoidal, strongly reticulate or rugose. The exocarp cells are isodiametric and convex to papillate. Also common are cylindrical nutlets, often with longitudinal ridging and papillate exocarp cells. Surface pitting and concave exocarp cells are rare. A cladistic analysis of nutlet characters suggests both Hemigenia and Microcorys are polyphyletic, and Microcorys paraphyletic with respect to Westringia Sm. Notwithstanding that, the infrageneric classification of Hemigenia was largely supported, while in Microcorys, there was support for sect. Hemigenioides, but sects Anisandra and Microcorys were not resolved as distinct.     

Fruit differentiation, palynology, and systematics in theScabiosa group of genera andPseudoscabiosa (Dipsacaceae)

Fruits ofDipsacaceae are single-seeded, have bristle-shaped calyx segments and are tightly enclosed by four fused bracts forming an epicalyx. Comparative morphological and anatomical studies reveal a great diversity of epicalyx and calyx, often relevant to fruit dispersal. The present contribution deals with theScabiosa group of genera, the core of theScabioseae tribe. Most of its taxa develop a diaphragma from a meristem on the inside of the epicalyx. This diaphragma, together with the lower part of the epicalyx encloses the fruit proper, whereas the upper parts form a so-called epi-diaphragma (ed) and a ± hyaline corona. Differences of the epicalyx with respect to the size and position of the ed, elaboration of the corona, origin of pits (=foveoles) and other morphological and anatomical specializations can be demonstrated. Together with palynological and karyological data these new facts support an improved concept of relationships and systematics for the taxa studied:Scabiosa sect.Scabiosa and sect.Cyrtostemma are closely related and should be united to form the genusScabiosa s. str.;Pycnocomon can be maintained as an independent genus, sister toScabiosa sect.Trochocephalus which then has to be treated as a genus,Lomelosia. In contrast, the following genera have to be included inLomelosia:Tremastelma asLomelosia sect.Callistemma, andScabiosiopsis as part ofLomelosia sect.Lomelosia. Pseudoscabiosa deviates in so many features that it has to be excluded from the redefinedScabioseae s. str.     

《广西植物志》秋海棠属(Begonia L.)增订

秋海棠属(Begonia L.)隶属于葫芦目(Cucurbitales)秋海棠科(Begoniaceae),有1 800余种,是世界第六大属,主要分布于美洲、非洲和亚洲的热带和亚热带地区,具有极高的观赏价值,亦可作为药用、食用、饮料和饲料等。我国有秋海棠属植物近300种,主要分布于我国的云南和广西等长江以南地区,其中云南有秋海棠属植物100余种。近年来,随着广西地区秋海棠属植物野外调查的不断深入,陆续发现并发表了大量秋海棠属新类群,使得广西产秋海棠属植物数目已超出1991年《广西植物志》(第一卷)收录的19种,因此亟需对发表于不同期刊中的类群进行系统地整理和汇总。鉴于此,该文通过查阅文献资料和考证标本信息,整理并汇总了广西产秋海棠属植物共84种,包括2个变种、11个亚种和46个特有种,新增的65种为《广西植物志》(第一卷)中未收录的类群,并补充了这些类群的特征集要、凭证标本信息和属下分组概况,为今后开展秋海棠属植物的分类修订以及系统学、进化生物学和保护生物学等研究提供了参考依据。     

中国赤车属新分类

(1)根据该文中赤车属与楼梯草属各种形态学的区别,认为赤车属较为原始,在楼梯草属之前自冷水花属衍生而出。(2)将赤车属划分为5组;将C.B.Robinson描述的Elatostemoides属降为组级群,作为赤车属中接近原始群全缘赤车组sect.Pileoides的一个组处理,此外还描述了一个与sect.Elatostemoides近缘的1新组,羽脉赤车组sect.Leiolaena,并根据叶、叶的脉序、雌花序、雌花被片和瘦果等器官的演化趋势给出显示赤车属属下诸群间亲缘关系的系统发育树。(3)对中国赤车属的属下分类群进行了修订,共承认32种1亚种7变种,给出检索表、描述及多幅插图,并将它们划分为4组、2亚组和9系,其中除sect.Leiolaena之外,还新描述了2亚组、5系、2种和3变种;在2003年被归并为同物异名的长柄赤车、曲毛赤车、波缘赤车、小赤车和富宁赤车的种级地位得到恢复。此外,还写出了中国赤车属的分类学简史和地理分布。     

Floral biology of Nuphar pumila (Timm) DC. (Nymphaeaceae) in China

Anthesis in individual flowers of Nuphar pumila (Timm) DC. occurs for four consecutive days. First-day flowers are protogynous and functionally female. Flowers can open completely on the first day of anthesis. This contrasts with all previous reports, which state that first-day flowers of Nuphar are characterized by partial expansion of the calyx, leaving a distal small triangular opening just above the stigmatic disc. Flowers close completely on the first night of anthesis and remain partially open on the subsequent three nights. During the entire anthesis period the stigmas emit a sweet, fruity odor and the petal nectaries produce visible nectar drops. The stigma of N. pumila is secretory and unicellular-papillate. Pollen grains are monosulcate with long spines. Our observations on the mating system of N. pumila indicate that neither asexual seed production nor spontaneous self-pollination occurs. Cross-pollination of second-, third- and fourth-day flowers produced few seeds. Flowers of N. pumila were mainly pollinated by sweat bees, with flies playing a minor pollination role. No beetle visits were observed. Our insect-pollination observations substantiate the view that the relative contribution of flies, bees, and beetles to pollination in a single Nuphar population depends on two factors: the relative abundance of the insects, and presence of alternative food sources.     

A phenetic study of Cassia sensu lato (Leguminosae-Caesalpinioideae: Cassieae: Cassiinae) in Thailand

Cassia L. sensu lato, a large heterogeneous genus of flowering plants, occurs naturally in the tropics around the world. Recent works based on floral morphology have supported a division of this genus into three genera, namely Cassia L. s. str., Chamaecrista Moench and Senna Mill. In order to investigate this new classification, 508 specimens of 18 taxa of the genus Cassia s.l. grown in Thailand were analyzed using cluster analysis and canonical discriminant analysis. The total 32 vegetative and reproductive morphological characters were employed in these analyses. In cluster analysis, Cassia s.l. can be separated into four groups, respectively viz. Chamaecrista, Senna alata, Senna and Cassia s /str. The four-cluster grouping is discussed. From a canonical discriminant analysis using the four-cluster grouping as a priori groups, it can be concluded that Cassia s. str., Senna, and Chamaecrista are indeed distinct taxa. The three most important characters that separate the three genera are filament length, fruit length, and ovary stalk length. These quantitative characters, together with some qualitative characters, were useful in constructing an identification key to these genera. Among the three genera, it was also found that Senna is rather a heterogeneous taxon. The difference between the studied species was discussed.     

A molecular phylogeny of Hebeloma species from Europe

In order to widen the scope of existing phylogenies of the ectomycorrhizal agaric genus Hebeloma a total of 53 new rDNA ITS sequences from that genus was generated, augmented by sequences retrieved from GenBank, and analysed using Bayesian, strict consensus and neighbour joining methods. The lignicolous Hebelomina neerlandica, Gymnopilus penetrans, and two species of Galerina served as outgroup taxa. Anamika indica, as well as representatives of the genera Hymenogaster and Naucoria, were included to test the monophyly of Hebeloma, which is confirmed by the results. Hebeloma, Naucoria, Hymenogaster and Anamika indica cluster in a strongly supported monophyletic hebelomatoid clade. All trees largely reflect the current infrageneric classification within Hebeloma, and divide the genus into mostly well-supported monophyletic groups surrounding H. crustuliniforme, H. velutipes, H. sacchariolens, H. sinapizans, and H. radicosum, with H. sarcophyllum being shown at an independent position; however this is not well supported. The section Indusiata divides with strong support into three groups, the position of the pleurocystidiate Hebeloma cistophilum suggests the possible existence of a third subsection within sect. Indusiata. Subsection Sacchariolentia is raised to the rank of section.     

Pollen morphology of Stachys (Lamiaceae) in Iran and its systematic implication

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.     

Comparative floral and vegetative differentiation between two European Aquilegia taxa along a narrow contact zone

As a first step in determining the identity and relative importance of the evolutionary forces promoting the speciation process in two closely related European taxa of Aquilegia, we investigated the levels of morphological variation in floral and vegetative characters over the narrow region where their ranges enter into contact, and evaluate the relative importance of both types of traits in their differentiation. A total of 12 floral and ten vegetative characters were measured on 375 plants belonging to seven A. vulgaris populations and six A. pyrenaica subsp. cazorlensis populations located in southeastern Spain. Floral and vegetative morphological differentiation occur between taxa and among populations within taxa, but only vegetative characters (particularly plant height and leaf petiolule length) contribute significantly to the discrimination between taxa. Differentiation among populations within taxa is mostly explained by variation in floral traits. Consequently, morphological divergence between the two taxa cannot be interpreted as an extension of among-population differences occurring within taxa. Multivariate vegetative, but not floral, similarity between populations could be predicted from geographical distance. Moreover, the key role of certain vegetative traits in the differentiation of A. vulgaris and A. p. cazorlensis could possibly be attributable to the contrasting habitat requirements and stress tolerance strategies of the two taxa. These preliminary findings seem to disagree with the currently established view of the radiation process in the genus Aquilegia in North America, where the differentiation of floral traits seems to have played a more important role.     

Biosystematic studies inTulipa sect.Eriostemones (Liliaceae)

Evolutionary relationships and infra- and interspecific variability within the species ofTulipa sect.Eriostemones Boissier were studied. Measurements of 35 morphological characters were used for principal component and canonical variate analyses. Crossing experiments and chromosome counts were carried out. Two axes of morphological variation were encountered within the principal component plot. Variation of subsect.Biflores was traced along one axis, the species of subsect.Australes as well asSaxatiles were found along the other axis. The nomenclature of names of subsect.Biflores was considered on the basis of their geography, specific morphological characteristics and polyploidy. Conspecificity of names of all species was tested based on both morphology and crossability.     

Variation in wild populations of annual beet (Beta,Chenopodiaceae)

Among the morphological variation in wild annual populations of sect.Beta only tepal characters show a geographic pattern, and hence can be used to distinguish different taxa. Two morphological types correspond with taxa already described:B. macrocarpa (incl.B. bourgaei) has a Macaronesian and W. to E. Mediterranean andB. adanensis an E. Mediterranean distribution area. A third type in the Aegean region is not well known yet and possibly has to be included inB. macrocarpa. Both diploid and tetraploid (x = 9) cytotypes are found withinB. macrocarpa, the latter exclusively on the Canary Islands.     

Phylogeny of Neolitsea (Lauraceae) inferred from Bayesian analysis of nrDNA ITS and ETS sequences

ITS and ETS-based sequence analyses of 29 Neolitsea, six Actinodaphne and five outgroup `core' Laureae taxa show that Neolitsea is monophyletic with two large subclades, whereas most of the sampled Actinodaphne are paraphyletic below it. Inflorescence features appear to be among the more reliable morphological characters for explaining relationships between Neolitsea and other genera within the `core' Laureae, with the Neolitsea/Actinodaphne clade defined by inflorescences lacking vegetative terminal buds in the main axis. Although the relationships within Neolitsea are still poorly resolved, there is enough structure to suggest that the genus seems to divide into two groups based on fruit shape: elliptic or ovoid, versus globose, although more evidence (both molecular and morphological) and wider taxon sampling are required to confirm this.     

Mating studies, new species, and new reports of Marasmius from northern Thailand

The mating systems operating in seven species of Marasmius collected recently in northern Thailand were determined. Marasmius cremeus belonging to sect. Sicci subsect. Siccini ser. Leonini, and Marasmius straminiceps belonging to sect. Marasmius subsect. Sicciformes are described as new species. Five members of sect. Marasmius were tetrapolar (bifactorial), viz. M. apatelius, M. guyanensis, M. nigrobrunneus, M. ruforotula, and M. straminiceps. Two members of sect. Sicci were bipolar (unifactorial), viz., M. corneri (syn. M. incarnatus) and M. cremeus. Our data support the hypothesis that the mating system is consistent within infrageneric taxa. The seven species that are herein described, illustrated, and compared with phenetically similar species represent the first reports for this genus in Thailand.     

Species relationships and taxonomy inTulipa subg.Tulipa (Liliaceae)

Principal component and canonical variate analyses were used to analyse the variation of 34 morphological characters measured in 98 accessions ofTulipa subg.Tulipa, formerly known as sect.Leiostemones. In addition 43 accessions were analysed cytogenetically by means of C-banding methods. Based on the encountered variation and on geographical distribution data 30 species in five sections were recognized, of which the synonymy is stated. Seven new series in two different sections have been designated.     

Detection of species within the Xerocomus subtomentosus complex in Europe using rDNA–ITS sequences

Identification of species within the boletoid genus Xerocomus has relied heavily upon the macromorphological features of the basidiomes. However, the phenotypic plasticity of these features has resulted in considerable confusion over the delimitation of taxa. In this study, we examined collections attributed to the X. subtomentosus complex in Europe using morphological and rDNA–ITS sequence data. In total, 45 European collections from a wide range of geographical areas and ecological conditions were included in the study. In spite of detecting considerable genetic variation, even within individual basidiomes of X. subtomentosus, molecular data, spore size, flesh colour, and the colour of the basal mycelium allow for the recognition of four distinct taxa: two correspond to X. subtomentosus (13 collections) and X. ferrugineus (20); one X. chrysonemus sp. nov. (10), to date only found in the UK, is described as new; and the existence of another taxon (two; Italy and UK) is noted but left undescribed owing to lack of material. Eight collections from North America were also included in the study, from which two taxa with a close affinity to X. ferrugineus were recognised.     

Morphological and molecular comparisons ofCampsis grandiflora andC. radicans (Bignoniaceae), an eastern Asian and eastern North American vicariad species pair

Morphological and molecular comparisons were made forCampsis grandiflora (Thunb.)K. Schumann (Bignoniaceae) from eastern Asia andC. radicans (L.)Seemann from eastern North America. Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases. The cpDNA divergence between the two vicariad species was 2.44%, which is the highest reported among North Temperate disjunct taxa and one of the highest reported for infrageneric taxa. Detailed morphological comparisons also suggest a high level of divergence. Cluster analyses based on 22 morphological characters and 39 OTUs revealed two distinct groups corresponding with the two species. The average taxonomic distance between the two species was 1.806. Analyses of variance (ANOVA) revealed that 12 of the 18 quantitative characters differed significantly ( 0.01) betweenC. grandiflora andC. radicans. Divergence time based on cpDNA data was estimated as 24.4 million years. The Bering land bridge hypothesis was favored over the North Atlantic land bridge hypothesis based on the estimated divergence time and the geological history of the North Temperate region. The high levels of morphological and cpDNA divergence are not consistent with morphological stasis, which has been proposed as a common mode of evolution for North Temperate disjunct taxa.