A chemotaxonomic survey of flavonoids and simple phenols in the leaves of Gaultheria L. and related genera (Ericaceae)

MIDDLETON, D. J., 1992. A chemotaxonomic survey of flavonoids and simple phenols in the leaves of Gaultheria L. and related genera (Ericaceae). Species of Gaultheria and Leucothoe have been screened for a number of flavonoids and simple phenols. Many of these, particularly the American species, are surveyed for the first time. Extensive sampling has also been carried out within some of the species of Gaultheria. Together with previously published data the results are discussed in relation to the position of Gaultheria in the Ericaceae and the infrageneric classification of Gaultheria. The infraspecific variation is commented on.     

Infrageneric classification of the genus Gaultheria L. (Ericaceae)

The genus Gaultheria has been classified into ten sections (one with two subsections) and 22 series of which five are new combinations and 21 are new taxa. Seven of these sections are composed primarily of solitary flowered species and the other three sections of racemose species. About half of the species of the genus are included in section Brossaea. A summary of the classification is here presented.     

果皮和种皮微形态特征在杜鹃花属系统学研究中的应用

采用扫描电子显微镜对国产杜鹃花属Rhododendron6个亚属的代表种和近缘类群杜香属Ledum杜香L.palustre的果皮和种皮微形态特征进行观察,对杜鹃花属植物果皮微形态特征进行了系统描述,并通过比较现存杜鹃花属植物和种子化石,新发现一些居间的种子类型。结果表明,果皮和种皮微形态特征具有一定的系统学意义。叶状苞亚属subgen.Therorhodion的叶状苞杜鹃R.redowskianum的果实有短而疏的刺毛,无气孔,种子为无翅类,扁平,外围轮廓长椭圆形。杜鹃亚属subgen.Rhododendron植物果皮为百合花杜鹃型,具有鳞片,气孔器散生于指状突起之间,与叶表皮的微形态特征一致,其种子为百合花杜鹃型,表面具宽而浅的沟,呈脑纹状,有别于无鳞类杜鹃花。常绿杜鹃亚属subgen.Hymenanthes果皮为云锦杜鹃型,其角质层表面不规则,无表皮毛,偶见气孔;种子为云锦杜鹃型。映山红亚属subgen.Tsutsusi果皮为岭南杜鹃型,密生长表皮毛,角质层致密;种子为南边杜鹃型和岭南杜鹃型。微形态特征与“常绿杜鹃亚属和(国产)映山红亚属均为内部支持率很高的单系类群”的分子系统发育研究结果一致。马银花亚属subgen.Azaleastrum的马银花组sect.Azaleastrum和长蕊杜鹃组sect.Choniastrum微形态区别明显,支持各自为独立的单系类群。羊踯躅亚属subgen.Pentanthera的羊踯躅R.molle果皮特征明显,可与其他亚属区分,但种子类型更接近常绿杜鹃亚属。本研究结合分子系统发育资料和叶表皮微形态特征讨论了一些近缘类群的系统发育关系;研究结果支持将腺萼马银花处理为独立的种。     

杜鹃花科白珠树属分子系统学和生物地理学研究进展

白珠树属(Gaultheria) 在杜鹃花科(Ericaceae)系统演化中占有十分重要的地位, 其系统位置和演化关系一直备受争议。最近的分子系统学研究认为, 白珠树属已经不再属于传统上的越橘亚科(Vaccinioideae)的綟木族(Andromedeae), 而是与一些相关属组成了白珠树族(Gaultherieae)。对白珠树属产于美洲的类群和相关类群的分子系统学的初步研究则表明, 该属与Diplycosia、Tepuia和Pernettya等属(均为“常绿类群”)关系密切, 可能应将这几个属并入到白珠树属中, 但其属下分类系统关系还需要对产于亚洲的类群进行深入的研究后才能确定。白珠树属与其近缘属的进化历史和生物地理学关系较为复杂, 与杜鹃花科其他大多数属不同, 白珠树属为典型的环太平洋分布。关于白珠树属的起源问题存在两种不同的推测: 一种观点认为该属起源于南半球的冈瓦纳古陆; 另一种观点则认为其起源于北半球的劳亚古大陆。本文概述了近年来白珠树属的分子系统学和生物地理学研究进展, 并对该属尚存的一些问题进行了讨论。     

杜鹃花科白珠树属分子系统学和生物地理学研究进展

白珠树属(Gaultheria)在杜鹃花科(Ericaceae)系统演化中占有十分重要的地位,其系统位置和演化关系一直备受争议.最近的分子系统学研究认为,白珠树属已经不再属于传统上的越橘亚科(Vaccinioideae)的綟木族(Andromedeae),而是与一些相关属组成了白珠树族(Gaultherieae).对白珠树属产于美洲的类群和相关类群的分子系统学的初步研究则表明,该属与Diplycosia、Tepuia和Pernettya等属(均为"常绿类群")关系密切,可能应将这几个属并入到白珠树属中,但其属下分类系统关系还需要对产于亚洲的类群进行深入的研究后才能确定.白珠树属与其近缘属的进化历史和生物地理学关系较为复杂,与杜鹃花科其他大多数属不同,白珠树属为典型的环太平洋分布.关于白珠树属的起源问题存在两种不同的推测:一种观点认为该属起源于南半球的冈瓦纳古陆;另一种观点则认为其起源于北半球的劳亚古大陆.本文概述了近年来白珠树属的分子系统学和生物地理学研究进展,并对该属尚存的一些问题进行了讨论.     

A chemosystematic study of the genus Gomphostemma and related genera (Lamiaceae)

Species in the genera Gomphostemma, Chelonopsis and Bostrychanthera were systematically studied with reference to their flavonoid and phenolic acid compounds in order to investigate whether the profiles of these compounds would support a classification of the genus and related genera based on morphological characters. Thirty-five flavonoid glycosides, eight phenolic acids and derivatives were identified by LC-UV-MS/MS analysis of aqueous 80% MeOH extracts on the basis of their UV and mass spectra, retention times and comparison with in-house library. The occurrence of individual compounds was not particularly informative in Gomphostemma, although the overall chemical profile supported G. subgen. Pogosiphon and vicenin-2 was a characteristic component of Gomphostemma leptodon and Gomphostemma curtisii. In contrast, the flavonoids and phenolic acids of Chelonopsis were informative at infrageneric level. Glycosides of 6-substituted flavones were well represented in Ch. subgen. Aequidens, including Ch. forrestii, Ch. rosea, Ch. odontochila, Ch. lichiangensis and C. giraldii. A dicaffeoylquinic acid was produced in Ch. subgen. Chelonopsis, (for example, in Ch. longipes and Ch. Moschata), but absent from Ch. subgen. Aequidens. The same dicaffeoylquinic acid was also found in the genus Bostrychanthera and suggests a close relationship with Ch. subgen. Chelonopsis, in agreement with a recent DNA based phylogeny. There is correlation between trichome type and phenolic acid compound distribution in Chelonopsis, but this is not observed in Gomphostemma.     

Consistent and variable leaf anatomical characters in Carex (Cyperaceae)

The anatomy and morphology of leaves in Carex have the potential to be taxonomically useful. However, studies on the variability of leaf characteristics in the genus are sparse. Researchers therefore risk using leaf anatomical characters without the knowledge of whether they are consistent in a species. We examined 22 qualitative and seven quantitative leaf anatomy characters from transverse leaf sections to test their consistency across 11 Carex spp. The characters were clearly described and primarily microscopic. Some characters were found to exhibit high levels of intraspecific variation, whereas other characters exhibited high levels of consistency in a species, including the shape of the leaf section, the density of papillae and the size of epidermal cells. Caution must be applied when choosing leaf anatomy to delimit taxa because of the intraspecific variability found in some characters, but sufficient numbers of invariant characters exist to provide useful taxonomic separation. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 371–384.     

DNA barcoding of Gaultheria L.in China (Ericaceae: Vaccinioideae)

Four DNA barcoding loci,chloroplast loci rbcL,matK,trnH-psbA,and nuclear locus internal transcribed spacer (ITS),were tested for the accurate discrimination of the Chinese species of Gaultheria by using intraspecific and interspecific pairwise P-distance,Wilcoxon signed rank test,and tree-based analyses.This study included 186 individuals from 89 populations representing 30 species.For all individuals,single locus markers showed high levels of sequencing universality but were ineffective for species resolvability.Polymerase chain reaction amplification and sequencing were successful for all four loci.Both ITS and matK showed significantly higher levels of interspecific species delimitation than rbcL and trnH-psbA.A combination ofmatK and ITS was the most efficient DNA barcode among all studied regions,however,they do not represent an appropriate candidate barcode for Chinese Gaultheria,by which only 11 out of 30 species can be separated.Loci rbcL,matK,and trnH-psbA,which were recently proposed as universal plant barcodes,have a very poor capacity for species separation for Chinese Gaultheria.DNA barcodes may be reliable tools to identify the evolutionary units of this group,so further studies are needed to develop more efficient DNA barcodes for Gaultheria and other genera with complicated evolutionary histories.     

DNA barcoding of Gaultheria L. in China (Ericaceae: Vaccinioideae)

Abstract Four DNA barcoding loci, chloroplast loci rbcL, matK, trnH‐psbA, and nuclear locus internal transcribed spacer (ITS), were tested for the accurate discrimination of the Chinese species of Gaultheria by using intraspecific and interspecific pairwise P‐distance, Wilcoxon signed rank test, and tree‐based analyses. This study included 186 individuals from 89 populations representing 30 species. For all individuals, single locus markers showed high levels of sequencing universality but were ineffective for species resolvability. Polymerase chain reaction amplification and sequencing were successful for all four loci. Both ITS and matK showed significantly higher levels of interspecific species delimitation than rbcL and trnH‐psbA. A combination of matK and ITS was the most efficient DNA barcode among all studied regions, however, they do not represent an appropriate candidate barcode for Chinese Gaultheria, by which only 11 out of 30 species can be separated. Loci rbcL, matK, and trnH‐psbA, which were recently proposed as universal plant barcodes, have a very poor capacity for species separation for Chinese Gaultheria. DNA barcodes may be reliable tools to identify the evolutionary units of this group, so further studies are needed to develop more efficient DNA barcodes for Gaultheria and other genera with complicated evolutionary histories.     

The systematic significance of morphological and anatomical variation in fruits of Crotalaria and related genera of tribe Crotalarieae (Fabaceae)

The phylogenetic and taxonomic significance of morphological and anatomical trends in fruits of tribe Crotalarieae has been evaluated, with emphasis on the genus Crotalaria and its seemingly distinctive, inflated and balloon‐shaped pods. In addition to the normal explosive dehiscence as a means of dispersal, several genera (including Crotalaria) show independent evolution of modifications apparently adapted for dispersal by wind, water and gravity. Transverse sections were made of mature pods of 142 species from the 12 currently recognized genera of Crotalarieae. The taxa differ in the orientation of the fibres (related to dehiscence or non‐dehiscence), the overall thickness of the fruit wall, the relative proportions of the pericarp layers, the degree of lignification and the presence or absence of trichomes. Three basic pericarp types can be distinguished: type I, with one, two or three zones of various numbers of cell layers of fibres (almost all genera); type II, with a single cell layer of fibres (only in Rothia, Robynsiophyton, Lebeckia and Lotononis sections Listia and Leobordea); and type III, with one zone of several cell layers of gelatinous fibres and multicellular trichomes associated with the endocarp (only in some species of Calobota and Wiborgiella). Considerable variation was encountered in the tribe, but Crotalaria appears to be rather uniform, with type I predominating. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 84–106.     

Alcantarea (Bromeliaceae) leaf anatomical characterization and its systematic implications

Alcantarea (Bromeliaceae) has 26 species that are endemic to eastern Brazil, occurring mainly on gneiss–granitic rock outcrops (‘inselbergs’). Alcantarea has great ornamental potential and several species are cultivated in gardens. Limited data is available in the literature regarding the leaf anatomical features of the genus, though it has been shown that it may provide valuable information for characterizing of Bromeliaceae taxa. In the present work, we employed leaf anatomy to better characterize the genus and understand its radiation into harsh environments, such as inselbergs. We also searched for characteristics potentially useful in phylogenetic analyses and in delimiting Alcantarea and Vriesea. The anatomical features of the leaves, observed for various Alcantarea species, are in accordance with the general pattern shown by other Bromeliaceae members. However, some features are notable for their importance for sustaining life on rock outcrops, such as: small epidermal thick‐walled cells, uneven sinuous epidermal walls, hypodermis often differentiated into lignified layers with thick‐walled cells, aquiferous hypodermis bearing collapsible cells, and the presence of well developed epicuticular stratum. Alcantarea leaves tend to show different shapes in the spongy parenchyma, and have chlorenchymatous palisade parenchyma arranged in more well‐defined arches, when compared to Vriesea species from the same habitat.     

Plastome characteristics and species identification of Chinese medicinal wintergreens (Gaultheria,Ericaceae)

Wintergreen oil is a folk medicine widely used in foods, pesticides, cosmetics and drugs. In China, nine out of 47 species within Gaultheria (Ericaceae) are traditionally used as Chinese medicinal wintergreens; however, phylogenetic approaches currently used to discriminating these species remain unsatisfactory. In this study, we sequenced and characterized plastomes from nine Chinese wintergreen species and identified candidate DNA barcoding regions for Gaultheria. Each Gaultheria plastome contained 110 unique genes (76 protein-coding, 30 tRNA, and four rRNA genes). Duplication of trnfM, rps14, and rpl23 genes were detected, while all plastomes lacked ycf1 and ycf2 genes. Gaultheria plastomes shared substantially contracted SSC regions that contained only the ndhF gene. Moreover, plastomes of Gaultheria leucocarpa var. yunnanensis contained an inversion in the LSC region and an IR expansion to cover the ndhF gene. Multiple rearrangement events apparently occurred between the Gaultheria plastomes and those from several previously reported families in Ericales. Our phylogenetic reconstruction using 42 plastomes revealed well-supported relationships within all nine Gaultheria species. Additionally, seven mutational hotspot regions were identified as potential DNA barcodes for Chinese medicinal wintergreens. Our study is the first to generate complete plastomes and describe the structural variations of the complicated genus Gaultheria. In addition, our findings provide important resources for identification of Chinese medicinal wintergreens.     

中国伞形科天胡荽亚科果实解剖特征及其系统学意义

经对中国伞形科天胡荽亚科 (Hydrocotyloideae)中天胡荽属 (HydrocotyleL .)积雪草属 (CentellaL .)和马蹄芹属(DickinsiaFranch .) 10种植物果实解剖结构的观察比较 ,发现 3属的果实横切面以合生面为基准可以分成 2个类型 ,即两侧压扁类型 (天胡荽属和积雪草属 )和背腹压扁类型 (马蹄芹属 )。除此之外 ,中果皮的外侧是否有色素块沉积、内侧是否有木化细胞层或厚壁细胞层以及最内层是否形成晶体细胞层 ,内果皮细胞的层数、排列方向以及有无棱槽油管等性状 ,可以作为区分 3属的重要解剖学特征。在此基础上 ,结合外部形态特征 ,讨论了该亚科及其内部各属的解剖学特征、属间区别及其演化关系。认为马蹄芹属应归属于Mulineae族 ,另外 2个属应归属于天胡荽族(Hydrocotyleae) ;3属中积雪草属原始而马蹄芹属进化 ;天胡荽亚科可能不是自然的类群 ,它的分类系统和演化地位有待深入研究。     

Generic limits in Polygonurn and related genera (Polygonaceae) on the basis of floral characters

In order to investigate the taxonomic value of a number of characters in the elucidation of relationships within the collective genus Polygonum L., and segregate genera Atraphaxis L., Calligonum L., Pteropyrum Jaub. & Spach., Oxygonum Burch., Fagopyrum Mill., Harpagocarpus Hutch. & Dandy and Polygonella Michx., 83 species of the tribe Polygoneae emend. Jaretzky were studied. The following floral characters were examined: stamen number, insertion and external morphology; structure and morphology of the floral nectaries; vasculature and surface morphology of the tepals.
Problems in floral morphology in Polygonaceae are discussed, together with relationships within the Family. The tribal division proposed by Haraldson (1978) is supported, but a new and different delimitation of genera is proposed. The present data support a division of Polygonum L. sensu lato and its related taxa into two tribes: Polygoneae (containing Polygonum L. sensu stricto, Fallopia Adans., Oxygonum Burch., Pteropyrum Jaub. & Sparh., Atraphaxis L. and Calligonum L.), and Persicarieae (containing Fagopyrum Mill., Harpagocarpus Hutch. & Dandy, Persicaria Mill. and Koenigia L.). The genus Reynoutria Houtt. is included as a section of Fallopia Adans.; Bislorta Mill., Aconogonon Reichenb. and Tovara Adans. are included within Persicaria Mill.
A number of new combinations are proposed in Persicaria and Fallopia .     

Leaf flavonoids as systematic characters in the genera Lavandula and Sabaudia

A comprehensive survey of the leaf flavonoids of the genus Lavandula and the related Sabaudia group was carried out using two-dimensional paper chromatography and high-performance liquid chromatography. The flavonoid patterns obtained were found to be systematically informative at the infrageneric level. Three main groupings were identified: the first containing sections Lavandula, Dentata and Stoechas characterised by the accumulation of flavone 7-glycosides; the second containing sections Pterostoechas, Subnuda and Chaetostachys characterised by the accumulation of 8-hydroxylated flavone 7-and 8-glycosides; the third encompassing the Sabaudia group and accumulating both flavone and 8-hydroxylated flavone 7- glycosides. Such a grouping of taxa is congruent with data from other disciplines, although it is not recognised in any present classifications. The taxonomic and evolutionary implications of the flavonoid data are discussed.     

Taxonomic separation of the genera Prosthechea and Encyclia (Laeliinae: Orchidaceae) using leaf and root anatomical features

Despite formal separation based on molecular and morphological evidence, the genera Prosthechea Knowles & Westc. and Encyclia Hook. have not been studied in terms of their vegetative anatomy. In this study we examine 16 Brazilian species of these genera. Additionally, one species of subtribe Laeliinae and another from subtribe Oncidiinae were studied in order to evaluate the taxonomic consistency of the anatomic characters observed. Except for Epidendrum crassifolium Lindl., all species possess a velamen differentiated into epivelamen and endovelamen. Endodermal cells with uniformly thickened walls, calcium oxalate raphides and cells with phi thickenings in the cortical region were observed in all species. Silica bodies, raphides and fibre bundles in the mesophyll were common in leaves of all species studied. By contrast, flavonoid crystals were observed in both roots and leaves of Prosthechea only. Encyclia is characterized by the presence of fibre bundles on the subepidermal layer, and a cuticle, which is clearly thicker than that of Prosthechea . These data strongly support the separation of Prosthechea and Encyclia .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 293–303.