Trichome anatomy and taxonomy of Saxifraga (Saxifragaceae)

Trichomes on flower pedicels and leaves were examined in 213 species of Saxifraga L. and in the monotypic, segregate genera Zahlbrucknera Reichenb., Saxifragella Engl., Saxifragodes D. M. Moore and Saxifragopsis Small. Pedicel trichomes were found to be better taxonomic characters than foliar ones. Six main trichome types were identified: i) multiseriate, glandular; ii) uniseriate, glandular; iii) sessile, multicellular, glandular; iv) multiseriate, eglandular; v) uniseriate, eglandular; vi) unicellular, eglandular. The different types were distributed (except in certain species) on pedicels as follows. Multiseriate, glandular types occur in four series of Saxifraga sect. Micranthes (Haw.) D. Don, part of sect. Gymnopera D. Don, and in sects Ciliatae Haw., Trachyphyllum Gaudin, Xanthizoon Griseb., Ligulatae Haw., Porophyllum Gaudin, Porphyrion Tausch and Irregulares Haw. They also occur in the genus Saxifragopsis. Uniseriate, glandular types occur in five series of Saxifraga sect. Micranthes , part of sect. Gymnopera and Irregulares , and in sects. Miscopetalum (Haw.) Sternb., Cymbalaria , Griseb. Tridactylites (Haw.) Engl., Saxifraga and Stoloniferae Haw. They also occur in the genus Zahlbrucknera. Eglandular, multicellular trichomes occur occasionally in Saxifraga sects Micranthes and Gymnopera , more frequently in sects Ciliatae and Ligulatae , and are ubiquitous in sects Xanthizoon, Porophyllum and Porphyrion. Eglandular, unicellular trichomes occur only in Saxifragopsis and Saxifragodes. Sessile, glandular trichomes occur in a group of species from Saxifraga sect. Stoloniferae. Saxifragella is glabrous. This taxonomic distribution of trichome types generally supports Engler's classification of the genus, and is compatible with other evidence from palynology, embryology and seed morphology. The taxonomic disposition of the few exceptional species is discussed.     

Molecular phylogenetics,character evolution and systematics of the genus Micranthes (Saxifragaceae)

With c. 85 species, the genus Micranthes is among the larger genera of the Saxifragaceae. It is only distantly related to the morphologically similar genus Saxifraga, in which it has frequently been included as Saxifraga section Micranthes. To study the molecular evolution of Micranthes, we analysed nuclear ribosomal (internal transcribed spacer, ITS) and plastid (trnL–trnF) DNA sequences in a comprehensive set of taxa comprising c. 75% of the species. The molecular phylogenetic tree from the combined dataset revealed eight well‐supported clades of Micranthes. These clades agree in part with previously acknowledged subsections or series of Saxifraga section Micranthes. As these eight groups can also be delineated morphologically, we suggest that they should be recognized as sections of Micranthes. New relationships were also detected for some species and species groups, e.g. section Davuricae sister to sections Intermediae and Merkianae, and M. micranthidifolia as a member of section Micranthes. Species proposed to be excluded from the genus Micranthes for morphological reasons were resolved in the molecular tree in Saxifraga. Many morphological characters surveyed were homoplasious to varying extents. Micromorphological characters support comparatively well the clades in the phylogenetic tree. An updated nomenclature and a taxonomic conspectus of sections and species of Micranthes are provided. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 47–66.     

Circumscription and phylogeny of Saxifraga sect. Ciliatae: Evidence from nrDNA ITS sequences

The largest section of the genus Saxifraga (Saxifragaceae), sect. Ciliatae, consists of 175 morphologically diverse species. This section is mainly distributed in the Qinghai-Tibetan Plateau and adjacent regions of southwest China and more than 80% of the total number of species are endemic to this region. It remains unknown whether this section is monophyletic and up to now no study has been conducted on the infra-sectional phylogeny. In this study, ITS sequences of the nuclear ribosomal DNA were firstly determined for 33 species mainly from this section and related sections. We further downloaded the corresponding sequences of the same DNA region for the other 22 species of Saxifraga and Mitella from GenBank. All sequences were together used to construct the phy-logenetic trees. The main implications of the phylogenetic analyses include: (1) sect. Ciliatae, as traditionally defined, constitutes as a monophyletic clade and its sister group is a well supported clade that includes species from 8 sections such as sect. Porphyrion, sect. Saxifraga and sect. Mesogyne; (2) three morphological subsections, i.e., subsect. Gemmiparae, subsect. Hirculoideae and subsect. Rosulares were tentatively recovered despite the relatively low statistic bootstrap support for the last one; however, subsect. Flagellares and subsect. Hemi-sphaericae were not recognized as separate entities, and nested within subsect. Gemmiparae; (3) subsect. Hircu-loideae and subsect. Rosulares clustered together as sister subclades while subsect. Gemmiparae diverged early. In addition, our results suggest that the paired variation of ITS sequences in sect. Ciliatae is relatively low between the sampled species in spite of their diverse morphology. It is suggested that such a scenario may mirror rapid speciation in this section that probably trigged by the uplifts of the Qinghai-Tibetan Plateau and the extensive selection pressure under the alpine environments.     

A monograph of Boykinia, Peltoboykinia, Bolandra and Suksdorfia (Saxifragaceae)

An evaluation of the limits and infra-generic taxonomy of the genus Boykinia and its allies is presented. In the past, Boykinia has been split into the following segregate genera: Boykinia and Telesonix (from North America) and Neoboykinia and Peltoboykinia (from Japan). The situation is complicated by other closely related genera, the limits of which also need clarification. Narrowly defined, they comprise Suksdorfia, Hemieva, Bolandra, Sullivantia and Jepsonia (all from North America) and Hieronymusia (from South America). This monograph draws on a variety of sources for information, including a morphological comparison of all taxa, using a wide range of both living and herbarium material, and observations on leaf anatomy, trichomes, pollen grains and seeds; reference is also made to our previously published work on flavonoids, chromosome numbers and breeding relationships. General ecological observations were made during field work in western North America. The results of these investigations confirm that the genera form a closely related group. Much of the evidence combines to suggest the following conclusions. Boykinia includes nine species in three sections: section Boykinia (B. aconitifolia , B. intermedia , B. lycoctonifolia , B. major , B. occidentalis and B. rotundifolia) , section Renifolium (B. ruhardsonii) and section Telesonix (B. jamesii and B. heucheriformis). Peltoboykinia (P. tellimoides and P. watanabei ) is recognized as a distinct genus allied to both Saxifraga and Boykinia. Suksdorfia , expanded to include Hemieva and Hieronymusia , has three species: S. violacea, S. alchemilloides and S. ranunculifolia. Bolandra is very closely related to Suksdorfia but is retained as a separate genus containing two species, B. californica and B, oregana. Sullivantia and Jepsonia are accepted as genera related to, but distinct from, Boykinia.     

An outline of a revised classification of Saxifraga L.

GORNALL, R. J., 1987. An outline of a revised classification of Saxifraga. A revised classification of Saxifraga is presented in which 15 sections, 19 subsections and 34 series are recognized. A total of 394 names are accounted for, including the following new taxa, combinations and names: series Hirculoideae (Engler & Irmscher) Gornall, stat. nov.; series Lychnitidae (Engler & Irmscher) Gornall, stat. nov.; series Nulanles (Engler & Irmscher) Gornall, stat. nov.; series Cinctae (H. Sm.) Gornall, stat. nov.; series Gemmiparae (Engler & Irmscher) Gornall, stat. nov.; series Spinulosae (C. B. Clarke) Gornall, stat. nov.; subsection Rosulares Gornall, nom. nov.; subsection Serpyllifoliae Gornall, subscct. nov.; subsection Hemisphaericae (Engler & Irmscher) Gornall, stat. nov.; section Merkianae (Engler & Irmscher) Gornall, stat. nov.; subsection Stellares (Engler & Irmscher) Gornall, stat. nov.; series Birostres Gornall, ser. nov.; series Melanocentrae (Engler & Irmscher) Gornall, stat. nov.; series Astasianlhes (Sternberg) Gornall, stat. nov.; subsection Micranthes (Haworth) Gornall, stat. nov.; series Aulaxis (Haworth) Gornall, stat. nov.; series Dermasea (Haworth) Gornall, stat. nov.; series Micranthes (Haworth) Gornall, stat. nov.; scries Aretioideae (Engler & Irmscher) Gornall, stat. nov.; series Juniperifoliae (Engler & Irmscher) Gornall, stat. nov.; series Lilacinae Gornall, ser. nov.; series Marginatae (Engler & Irmscher) Gornall, stat. nov.; series Squarrosae (Engler & Irmscher) Gornall, stat. nov.; series Rigidae (Engler & Irmscher) Gornall, stat. nov.; subsection Engleria (Siindermann) Gornall, stat. nov; series Subsessiliflorae Gornall, ser. nov.; series Oppositiofoliae (Hayek) Gornall, stat. nov.; series Tetrameridium (Engler) Gornall, stat. nov.; subsection Mutatae (Engler & Irmscher) Gornall, stat. nov.; subsection Florulentae (Engler & Irmscher) Gornall, stat. nov.; section Odontophyllae Gornall, sect, nov.; series Biternatae (Engler & Irmscher) Gornall, stat. nov.; subsection Triplinervium (Gaudin) Gornall, stat. nov.; series Arachnoideae (Engler & Irmscher) Gornall, stat. nov.; subsection Tridactylites (Haworth) Gornall, stat. nov.     

Flavonoid variation in some North Americna Saxifraga species

Eight species of Saxifraga representing sections Micranthes, Hirculus, Dactyloides and Xanthizoon were studied for their flavonoids (S. california, S. integrifolia, S. michauxii, S. ferruginea, S. eschscholtzii, S. hirculus, S. caespitosa and S. aizoides). The major compounds present in most species were kaempferol and quercetin monogluocosides and galactosides. Glucosides of kaempferol and quercetin predominate in the first four species listed, while galactosides of quercetin and myricetin are dominant in the lastthree. 3-O-Methyl- and 3,3′-di-O-methylquercetin were identified from S. californica and S. integrifolia. Saxifraga caespitosa synthesizes a complex mixture of O-methylated flavonols as well as a novel O-methylated dihydrokaempferol.Species pairs S. michauxii/S. ferruginea and S. californica/S.integrifolia exhibit close flavonoid similarity which may reflect their morphological similarities.     

Pollen morphology in the genus Saxifraga and its taxonomic significance

The pollen morphology of 105 species of Saxifraga has been investigated. Four major types of sexine pattern are recognized, namely reticulate, granular, striate and very finely striate with superimposed granules or verrucae. The striate pattern is found to be the most common and is divided into eight subtypes. The types and subtypes are related to the sections recognized in the genus and are generally found to support the existing classifications. The pollen-types also clarify some of the longstanding problems but leave others unresolved, and some exceptional patterns occur within otherwise well-defined sections.     

A taxonomic study on Saxifraga ser. Radioflorae (Saxifragaceae) in Japan,the Kuril Islands and Kamchatka

Plants of Saxifraga ser. Radioflorae (A. M. Johnson) Zhmylev, now commonly regarded as belonging to the genus Micranthes, including Saxifraga fusca Maxim. and Saxifraga purpurascens Kom. (Saxifragaceae), distributed from Kamchatka to southern Japan through the Kuril Islands, are here taxonomically revised on the basis of floral morphology. The results suggest that the plants can be divided into three groups: 1) those from Kamchatka and the northern Kurils, 2) those from Kunashiri to the Japanese archipelago, through Hokkaido and Honshu, and 3) those from the southern Kurils from Chirpoi to Etorofu Islands. Leaf characteristics and habitat preferences also correlate with these groupings. The first group is identified as Micranthes ohwii ( = Saxifraga purpurascens Kom.), and the second group as M. fusca. The third group is similar to M. fusca in floral proportions, but has petals with non‐revolute margins, larger floral discs and disc and filaments developing throughout anthesis. The third group may be derived from M. fusca, adapted to the coastal environment.     

matK and rbcL gene sequence data indicate that Saxifraga (Saxifragaceae) is polyphyletic

The large genus Saxifraga, which consists of ≈400 morphologically and cytologically diverse species, has long been considered taxonomically complex. Phylogenetic analysis of over 2500 bp of chloroplast sequence data derived from matK and rbcL was employed to examine relationships among sections of Saxifraga, the segregate genera Zahlbrucknera, Saxifragopsis, and Cascadia, and the relationships of these taxa to other Saxifragaceae sensu stricto. Phylogenetic trees resulting from separate analyses of the matK and rbcL sequences were highly congruent; phylogenetic analysis of a combined matK–rbcL data matrix was therefore also conducted. Our analyses indicate that Saxifraga is polyphyletic, comprising two well-differentiated clades. One clade, Saxifraga sensu stricto, is the sister to the remainder of the family and consists of Saxifraga sections Irregulares, Heterisia, Trachyphyllum, Cymbalaria, Mesogyne, Xanthizoon, Porphyrion, Ciliatae, Cotylea, Ligulatae, Saxifraga, and Gymnopera. With the exception of Gymnopera, the species-rich sections of this clade are monophyletic. Also part of this clade is the problematic Zahlbrucknera paradoxa, which is allied with members of section Saxifraga. A second major clade of Saxifraga species, Micranthes sensu lato, comprises the large section Micranthes, as well as the segregate genus Cascadia, and S. tolmiei of section Merkianae. This clade is allied with the Heuchera, Darmera, and Chrysosplenium-Peltoboykinia groups of genera. The segregate genus Saxifragopsis is only distantly related to species of Saxifraga, and is instead the sister to Astilbe. The monotypic Oresitrophe is confirmed as a member of the Darmera group of genera. These results suggest that the floral features used to define Saxifraga may simply be symplesiomorphic in these well-separated Saxifraga lineages. Furthermore, the enormous cytological diversity encompassed by Saxifraga likely represents two independent instances of extensive aneuploidy and polyploidy in Saxifragaceae.     

Phylogenetic relationships of the silver saxifrages (Saxifraga, sect. Ligulatae haworth): implications for the evolution of substrate specificity, life histories, and biogeography

The silver saxifrages (Saxifraga sect. Ligulatae Haworth; Saxifragaceae) exhibit remarkable variation of substrate specialization, with strictly calcicole to calcifuge species, as well as life histories which range from semelparity to iteroparity. They occur almost exclusively in the European mountain ranges and display high levels of endemism. Sequences from chloroplast and nuclear ribosomal DNA were obtained to resolve phylogenetic relationships among the silver saxifrages and related taxa and to gain insight into the evolution of substrate specificity, life history, and biogeography. The resulting phylogenies suggested that (1) Saxifraga sect. Ligulatae, as traditionally defined, does not constitute a monophyletic group; (2) lime-secreting hydathodes in calcifuge species apparently represent a secondary nonaptation; (3) semelparity evolved independently two or three times in the silver saxifrages and allied sections, possibly in response to climatic changes that occured during the Pleistocene; and (4) narrow endemics, for example S. cochlearis, likely evolved from the fragmentation of the widespread S. paniculata into refugial populations that became isolated during the glacial maxima of the Pleistocene.     

THE DISTRIBUTION OF ANTHOCHLOR FLORAL PIGMENTS IN NORTH AMERICAN COREOPSIS (COMPOSITAE): TAXONOMIC AND PHYLETIC INTERPRETATIONS

Anthochlors (chalcones and aurones) occur in floral tissues of all members of North American Coreopsis. The genus in North America consists of 46 species distributed in 11 sections, and a total of five chalcone aglycones are found in it. The chalcone butein occurs in all sections, and is the only anthochlor constituent in the closely related woody Mexican sections Electra and Anathysana. These two sections are viewed as primitive. The morphologically distinct but putatively unspecialized woody Mexican section Pseudo-Agarista sequesters both okanin and butein; the same two anthochlors also characterize two additional sections of herbaceous perennials, Silphidium and Tuckermannia. Sections Euleptosyne and Pugiopappus, comprised of annuals restricted primarily to California, exhibit the rare chalcone stillopsidin in addition to butein and okanin. The remaining four sections, Palmatae, Coreopsis, Calliopsis and Eublepharis are among the more specialized in North America, and are sometimes considered to represent a distinct phyletic line or two related lines. All or some of the members of these sections sequester the very rare chalcone lanceolatin in floral tissues in addition to okanin. Butein is also present in some or all species in each section, and isoliquiritigenin occurs rarely in two sections. Those sections considered more specialized have additional anthochlors relative to primitive sections, and any compound found in a more primitive section also occurs in a more advanced one. Thus, the trend in floral anthochlors in North American Coreopsis is toward increased number of compounds rather than reduction.     

Foliar anatomy ofPolygonum (Polygonaceae): Survey of epidermal and selected internal structures

Several anatomical characters in leaves were described, and their distribution determined, for 153 species ofPolygonum, mostly from herbarium specimens. Structures surveyed were epidermal (glandular and nonglandular trichomes, nodules, specialized parenchyma, stomatal apparatus) and internal (cavities, crystals, laticifer-like cells, nodules, subepidermal fibers). Cleared leaves were supplemented by resin-embedded sections and SEM preparations of selected species. No feature defines any taxonomic section, but some features occur only within one section. Laticifer-like cells, epidermal and internal nodules, resin cups, and unique epidermal and subepidermal cavities seem to be unknown elsewhere; other features (invaginated epidermal cells; enlarged crystal cells confined to paraveinal layer) are rare among angiosperms.     

不同类型人类活动干扰对河岸带外来植物群落的影响——以北京永定河为例

强烈的人类活动对自然生境的扰动促进了外来植物侵入河岸带,但不同类型的人类活动干扰对外来植物的影响有所不同。按照不同人类活动干扰类型设置了山峡段、平原段和城市段,通过比较河段间河岸带外来植物群落特征的差异,探讨不同人类活动干扰类型对河岸带外来植物群落的影响。结果表明:(1)永定河河岸带共有维管束植物27科72属101种,外来植物13科28属29种。外来种比例高达28.7%,其中82.4%的国外外来种来自美洲和亚洲。(2)不同河段间的外来植物群落的物种构成和优势度呈现出显著性差异。外来种比例由山峡段的20.9%,上升至平原段和城市段的30.2%和25.5%;而优势度由12.1%,分别上升至13.4%和17.5%。(3)不同河段间的外来植物群落生活型结构有显著性差异。多年生草本植物在山峡段比例最高,达到66.7%,在平原段和城市段较低,分别为46.2%和30.8%;而一年生草本植物在城市段比例最高,达到69.2%,在平原段和山峡段较低,分别为53.8%和33.3%。(4)平原段属于农业干扰类型,其河岸带外来植物主要以农业类杂草为主,如禾本科的假稻(Leersia japonica)和菊科的钻叶紫菀(Aster subulatu);城市段是城市干扰类型,外来植物中入侵种比例和优势度较高,典型入侵种为反枝苋(Amaranthus retroflexus)和牛筋草(Eleusine indica)等。北京永定河不同河段河岸带外来植物的种类构成特点,反映了河岸带外来植物受快速城市化、农业活动等不同人类活动干扰的影响呈逐渐扩大的趋势。     

The Classification and Distribution of Genus Deutzia in China

Discussed in the present paper are evolutionary trends of important morphological characters of Deutzia, systematic position of several closely related genera, geographical distribution and characteristics of floristic elements. Finally the classificatory revison of the genus is made and a key to species is given. As a result, evolutionary trends of the important morphological characters in Deutzia are suggested; petals from imbricate to valvate, stamens from indefinite to definite, filaments from edentate to dentate, ovary from half-inferior to inferior. Therefore, the section Neodeutzia with valvate corolla and infinite stamens with edentate filaments should be included in this genus. The sections Neodeutzia and Mesodeutizia seem to be better considered as primitive taxa, while the section Deutzia advanced one. There are 52 species of Deutzia in China , which are grouped into two sections ( Sect. Mesodeutzia, Sect. Deutzia), four subsections (Subsect. Deutzia, Subsect. Grandiflorae, Subsect. Stenosepalae, Subsect. Cymosae)and 17 series. Sixty- five species are so far recognised in the genus Deutzia. They are mainly distributed in E. Asia and disjunctively in N. America. China is therefore an area the richest in species of Deutzia in the world, making up 80% of the total species of this genus. The greatest concentration area is in Sichuan (23 species), Yunnan (21 species), Hubei (12 species) and Shanxi (10 species), this is boundary area between Sino-Himalayan subregion and Sino-Japanese subregion, where occur abundant species (including 39 endemic species) and diverse taxa (2 section and 4 subsection). Based on these facts it is proposed that the present center of distribution and differentiation of Deutzia be in the southern part of the Hengduan Mountains, the Qingling Range and C. China.     

Phylogenetics of Annona cherimola (Annonaceae) and some of its closest relatives

Annona cherimola is a woody perennial species in the Annonaceae family that produces edible fruits and has economic importance in several regions of the world with subtropical climates. Together with other 10‐12 species, A. cherimola belongs to the section Atta of the Annona genus with a center of origin in Central America and the Caribbean. Species of the section Atta produce soft skin ripe fruits with raised areoles bounded by recessed furrows. Annona cherimola is the only species of the section naturally found in the Andean region of South America. Currently, no information is available at the molecular level on the phylogenetic relationships of most of the species of Atta and closely related sections in Annona. In order to fill this gap, in this work a phylogenetic approach was performed using five coding and non‐coding plastid DNA regions, to determine the phylogenetic relationships between A. cherimola and other related species included in Atta and other sections of the genus. The results obtained support recent studies that demonstrated the likely Mesoamerican origin of A. cherimola based on biogeographical analysis with SSR markers, rather than the previously considered South American origin hypothesis. In addition, the species belonging to the Atta section did not show monophyly. Finally, A. cherimola and A. pruinosa seem to be phylogenetically close species and additional studies are needed to discern the relations between them.     

A new endemic species of Saxifraga L. from Spain

A recently erected species of Saxifraga (section Porphyrion ) endemic to the Picos de Europa (Cordillera Cantábrica) is described and illustrated. S. felineri P. Vargas resembles S. aretioides and S. ferdinandi-coburgi , although the habit and leaf shape are the main characters which support its identity. The taxonomic relationships of the three species are discussed, as well as aspects of the distribution and ecology of S. felineri.     

Phylogeny of Iris based on chloroplast matK gene and trnK intron sequence data

Phylogenetic analyses of 46 species of Iris, representing all subgenera and all sections except Regelia, Brevituba, and Monolepis, utilized matK gene and trnK intron sequence data. Sequence data show that Iris is paraphyletic because Belamcanda chinensis is resolved within the genus. The two largest subgenera, Iris and Limniris, are both resolved as polyphyletic. With the removal of section Hexapogon, subgenus Iris is weakly supported as monophyletic. Relationships within subgenus Limniris are more complex with the subgenus as currently circumscribed representing eight independent origins among the species included in this study. Several potential monophyletic groups are identified including subgenus Scorpiris, series Spuria (subgenus Limniris section Limniris), and a clade of section Limniris species from North America and Asia.     

Observations on the genus Pandanus in Madagascar

The sections of the genus Pandanus which occur in Madagascar are the main subject of this paper. There are 75 species in the island, of which all, or all but one, are endemic. They can be grouped into 12 sections, of which six are endemic, and one is a local endemic subsection. The other sections occur elsewhere, mainly in Africa, but two to the east. A key to the sections is given, and a discussion of field recognition characters is included. One new section and one new subsection are proposed. New lectotypifications for Sections Sussea and Vinsonia are suggested. The Madagascar species of Thouars are discussed. Finally a resume of some recent collections is presented.