Review: Taxonomy of Nitella (Charales, Charophyceae) based on comparative morphology of oospores and multiple DNA marker phylogeny using cultured material

The genus Nitella is the largest group in the Charales and has the highest diversity of vegetative and oospore morphologies. In his worldwide monograph on the Charales, R. D. Wood characterized the sections and species of Nitella mainly on the basis of vegetative morphology and treated oospore wall morphology as diagnostic at the infraspecific level. Therefore, many species of Nitella exhibiting distinct external morphology of the oospore wall (EMOW) were reduced to infraspecific rank and only 53 of 204 species previously described were recognized within Nitella . However, recent morphological and molecular phylogenetic studies have demonstrated the phylogenetic validity of using EMOW for diagnosing some species of Nitella . More recently, a scanning electron microscopy (SEM) study of internal morphology of the oospore wall (IMOW) and multiple DNA marker analyses using both nuclear and chloroplast gene sequences were conducted to improve our understanding of the taxonomy of Nitella at the species level, on the basis of cultured material of a large number of species. Multiple DNA marker analyses resolved detailed and robust phylogenetic relationships within the genus and demonstrated the taxonomic and phylogenetic significance of IMOW. In addition, they supported the taxonomic decisions based on differences in oospore morphology, suggesting that an integrated approach, involving both SEM studies of the EMOW and IMOW and multiple DNA marker analyses, is appropriate to address problems at lower taxonomic levels within the genus, as well as to construct a natural taxonomic system for the genus. In this paper, recent morphological and molecular phylogenetic studies are reviewed and recent improvements in taxonomy of Nitella are summarized. Moreover, the evolution of morphological features and phylogenetic relationships within Nitella are discussed, focusing especially on oospore morphology.     

TAXONOMIC REEXAMINATION OF 17 SPECIES OF NITELLA SUBGENUS TIEFFALLENIA (CHARALES,CHAROPHYCEAE) BASED ON INTERNAL MORPHOLOGY OF THE OOSPORE WALL AND MULTIPLE DNA MARKER SEQUENCES1

In an attempt to reconstruct the natural taxonomic system for Nitella, 17 species of Nitella subgenus Tieffallenia were reexamined using SEM observations of the internal morphology of the oospore wall (IMOW) and phylogenetic analyses of 4553 base pairs from multiple DNA markers (atpB, rbcL, psaB, and ITS‐5.8S rRNA genes). Our SEM observations identified three types of IMOW: homogeneous (HG), weakly spongy (W‐SG), and strongly spongy (S‐SG) types. Based on differences in the IMOW, species with reticulate or tuberculate oospore wall ornamentation in the external morphology of the oospore wall (EMOW) were subdivided into two distinct groups (characterized by the HG or S‐SG types of IMOW, respectively), which were robustly separated from each other in our molecular phylogenetic analyses. In our molecular phylogeny, the subgenus Tieffallenia consisted of four robust monophyletic groups—three clades of the HG type and a spongy (S‐SG and W‐SG) type clade—that were characterized by differences in the IMOW and EMOW. In addition, our SEM observations and sequence data verified the distinct status of five species (N. japonica Allen, N. oligospira A. Braun, N. vieillardii stat. nov., N. imperialis stat. nov., and N. morongii Allen) that R. D. Wood had assigned as infraspecific taxa. Moreover, our SEM observations of the IMOW also suggested that N. megaspora (J. Groves) Sakayama originally identified by LM includes at least two distinct species, characterized by W‐SG and S‐SG types of IMOW, respectively.     

Some New Taxa of Charophyta in China

The authors examined specimens collected from Beijing, Hebei Province and North-east China. Two new species, one variety and five new records of the Characeae from China are reported in this paper. They are Nitella nenjiangensis sp. nov., Chara vertillibracteata sp. nov., C. globularis Thuiller var. beijingensis var. nov., Nitella axillaris Braun, N. crispa Imah., N. gracillima Allan, N. pseudoflabellata var. imperialis T. F. A., Chara arru-densis Mendes.     

Some New Taxa and Records of Characeae in Hunan,China

In this paper, one new species, four new varieties and thirteen new records of Characeae from Hunan, China, are reported. They are Chara quadriscutulum, sp. nov., Nitella axilliformis var. biformis, var. nov., N. microcarpa var. unicarpa, var. nov., Chara braunii var. cylindrospora, var. nov., C. hydropitys var. hunanensis, var. nov., Nitella microcarpa Braun, N. leptoclada Braun, N. annandalei Pal, N. confervacea A. Braun, N. gracilliformis J. Groves, N. inversa Imahori, N. musashiensis Morioka, N. paucicostata T. F. Allen, N. acuminata var. gollmeriana (A. Br.) Zanev. et Wood, Tolypella boldii Sawa. Chara longifolia Robinson, C. pistianensis J. Vilhelm, C. grovesii Pal.     

ANNUAL GERMINATION WINFOW IN OOSPORES OF NITELLA FURCATA (CHAROPHYCEAE)1

Oospores of Nitella furcata subsp. megacarpa (Allen emend. Wood) were collected from an oospore bank in the sediments of Lake George, New York. Incubated at constant temperatures, all or nearly all of the oospores germinated when exposed to a brief pulse of red light when the annual window of germinability was open. The window seems related to the annual cycle of sediment temperatures. It is open in spring and closes wit the onset of a secondary dormacncy in the summer. Oospores in storage follow a parallel path if held at 18°C, a summer equivalent temperature; the window remains open indefinitely if the oospores are held at 4°C. Attention is drawn to the similarity if the cyclic window of germinability in seeds of summer annuals and oospores of N. furcata.     

Oospore dimensions and wall ornamentation patterns in Swedish charophytes

The ultrastructural features of oospore wall ornamentation patterns revealed by scanning electron microscopy are important taxonomic characters of the Charales. The present study shows inter-and intrapecific variations in 19 species of the genera Chara, Lamprohamnium, Nitella and Tolypella. This is the first time that the oospore wall ornamentation of Swedish Charales has been documented in detail. In the studied Chara species the ornamentation within species was variable, and partly overlapping between species, but only between closely related species. In contrast, the Nitella species showed distinct differences in ornamentation patterns between species, although the same patterns can be found in different species. This study presents for the first time SEM images of the species Chara rudis (pustular ornamentation), Nitella opaca (pitted ornamentation) and Nitella wahlbergiana (anastomosing network ornamentation). The ornamentation pattern in the Nordic species TV. wahlbergiana , supports its separation from Nitella mucronata which has a reticulate ornamentation. The relationship between length and width of the oospores is also of taxonomic significance. Nitella and Tolypella oospores are roundish, whereas those of Chara and Lamprothamnium are elongate. The extent to which environmental and genetic factors can affect oospore size and shape within a species are still unknown, but in the present study both ornamentation pattern and size provided evidence for the distinction between Chara globularis and Chara aspera. Some further taxonomic problems in the Charales are discussed in the light of the results of this study.     

轮藻植物在澳大利亚的重新发现之探密

澳大利亚轮藻植物研究历史悠久,迄今约有95种被描记。在Wood(1972)对澳大利亚轮藻植物的修订研究中,一些种因为标本遗失或已损坏而未包括在内,其中有的种近年来又被重新发现。本文重点介绍Nitella hookeri A．Br．var．arthroglochin A．Br．,Lychnothamnus barbatus(Meyen)Leonh．和Nitella partita Nordst．的发现过程及其产地生态环境,探讨了季节性湿地对轮藻植物的保护意义。N．hookeri var．arthroglochin早在1854年就发现于澳大利亚,并于1887年再次作为澳大利亚特有物种而被记录,然而从此销声匿迹,直至1989年才再度问世,它重新出现的地点为新南威尔士州高海拔花岗岩风化土壤地区的季节性溪流中。L．barbatus则为R．D．Wood首次发现于1960年,嗣后一再搜寻并无踪迹,直到1996年才于昆士兰东南地区亚热带阵发性溪流中被再次发现。昆士兰中部干旱地区1889年一度见到的N．partita,也是近一百年后才在新南威尔士州西北干旱地区的季节性湿地再次发现。     

Some New Taxa of Charophyta from Guangxi

Guangxi, Situated in the southern border area of China, is extremely rich in Charophytes. In this paper, however, reported are only new species and new varieties. Among the new taxa, Nitella haplodactyla has only one dactyl on the fertile branchlet, N. brachyclema and N. rarissima are provided with more than 4-celled dactyls, the fertile branchlet of N. pseudoflabellata var. tetradynana has tetradynamous rays, and Chara piniformis is characterized by thehaplostichous cortex, long spinecells, and rudimentary stipulodes.     

Different cytoplasmic calcium contents among three species of Characeae.

Internodal cells of three species of Characeae, Nitella flexilis, Nitella axilliformis and Chara corallina, were analyzed for the contents of Ca(2+ )and Mg(2+) in the cytoplasm. To avoid contamination of Ca(2+) from the cell wall and vacuole, the vacuolar sap was replaced with a sorbitol solution containing Sr(2+) by the vacuolar perfusion method after the cell had been treated with Sr(2+). No significant difference in the cytoplasmic content of Mg(2+) was found among three species of Characeae, but significant differences in the cytoplasmic content of Ca(2+) were observed among them. The cytoplasmic Ca(2+) content of N. flexilis was 2.0 times that of N. axilliformis and 3.3 times that of C. corallina. The cytoplasmic drop was furthermore separated into two fractions: a chloroplast-free fraction and a chloroplast fraction. In the chloroplast-free fraction the Ca(2+) content of N. flexilis was 2.3 times that of C. corallina and 2.0 times that of N. axilliformis, while the Mg(2+) content was the same among the three species. In the chloroplast fraction N. flexilis contained about seven times more Ca(2+) and about two times more Mg(2+) than C. corallina. The difference in the cytoplasmic Ca(2+ )content was discussed in relation to the difference in the capacity for the hydration-induced Ca(2+) release existing among the three species.     

Evidence for the Polyphyly of Haworthia (Asphodelaceae Subfamily Alooideae; Asparagales) Inferred from Nucleotide Sequences of rbcL, matK, ITS1 and Genomic Fingerprinting with ISSR-PCR

Abstract: Four molecular markers have been studied to examine the phylogenetic position of the South African plant genus Haworthia Duval within the succulent Asphodelaceae. Sequence data of the chloroplast genes mat K and rbc L were compared to the nuclear markers ITS1 and ISSR (Inter Simple Sequence Repeat) analysis. Both lines of molecular data, chloroplast and nuclear DNA, indicate that Haworthia is polyphyletic, forming two distinct clades. Most taxa previously combined as Haworthia subgenus Haworthia branch off early in the alooid chloroplast trees forming a strongly monophyletic group, whereas subgenus Hexangulares forms a polyphyletic assemblage comprising other alooid genera. The nuclear markers ITS1 and ISSR fingerprinting support the two groups as distinctly different, therefore confirming the division seen in chloroplast DNA. The practical implication is that the generic concept of Haworthia may have to be restricted to H. subgenus Haworthia or alternatively, that the groups of Haworthia be treated as infrageneric taxa within a broadened (Linnaean) concept of Aloe.     

中国丽藻属两新种

本文报道的是采自广西、山东的丽藻属植物两新种,即广西丽藻(Nitella guangxiensis)和山东丽藻(Nitella shandongensis),其特征分别描述如下: 广西丽藻新种图1:1—9 Nitella guangxiensis Y. J. Ling, G. Z. Deng et Z. Li, sp. nov. Planta monoica, claro-viridis, 13-19 cm alta. Caulis robustus,diametro 610-     

RAMARIA SUBGENERA RAMARIA AND LAETICOLORA IN YUNNAN

Three taxa of Ramaria subgenus Ramaria, and 18 of subg. Laeticolora are reported from Yunnan Province. Of these 13 are considered new to science. Only one taxon (R. formosa) is considered of widespread distribution, the others being known olay from the Indo-Asian subcontinent. Wide distribution patterns, based on phenetic similarity, seem to be at the infraspecific to species complex level, but only rarely contaxie. New taxa are as follows: subg. Ramaria: R. eryuanensis; subg. Laeticolora: R. brunneipes, R. distinctissima, R. ephemeroderma, R. hilaris, R. lacviformosoides, R. linearioides, R. linearis, R. nanispora, R. rubri-attenuipcs, R. sino-conjunctipes, R. rubricarnata var. laeta, R. botrytoides var. microspora, and R. neoformosa var. sinensis. Keys to Yunnan taxa of Ramaria subgenera Ramarla and Laeticolora are furnished.     

PHYLOGENETIC UTILITY OF rbc S SEQUENCES: AN EXAMPLE FROM ANTITHAMNION AND RELATED GENERA (CERAMIACEAE, RHODOPHYTA)

We report the potential phylogenetic utility of the small RUBISCO subunit ( rbc  S) sequences from a sampling of Antithamnion and related genera in ceramiacean algae. The size of rbc  S was 417 bp for all taxa examined. Analyses of the DNA sequence data indicated that pairwise divergences of rbc  S sequences were 3.3%–9.8% among species of Antithamnion , and ranged from 13.6% to 18.0% between Antithamnion and related genera. Phylogenetic analyses fully resolved relationships at the intrageneric level with statistical significance supported by high bootstrap values. Two subgenera of Antithamnion , Pteroton and Antithamnion , were clearly distinguished in the molecular tree. In the clade of subgenus Pteroton , A. aglandum was allied with A. callocladum and separated from A. nipponicum. In the pairwise distance comparison of sequence variation, Ceramium showed the greatest genetic distances among genera examined in the study. All phylogenetic trees generated by the maximum parsimony, neighbor joining, and maximum likelihood were completely congruent in topology with high confidence.     

Lectotypifications and notes on the application of names of Hieracium sect. Hieracium, sect. Bifida and sect. Vulgata (Asteraceae) based on material from the province of Dalarna, central Sweden

Lectotypes are designated for 174 names of taxa described based on material from the Swedish province of Dalarna and accepted to belong to Hieracium sect. Hieracium , sect. Bifida and sect. Vulgata . The taxonomic and nomenclatural implications, i.e. taxonomic synonyms and infraspecific taxa not worthy of taxonomic recognition, of the lectotypifications are discussed. Three new combinations, H. mundulifolium (Johanss.) T. Tyler, H. oligasterum (Johanss. & Sam.) T. Tyler and H. transtrandense T. Tyler, are proposed to accommodate the infraspecific taxa H. gilvocaniceps Johanss. var. mundulifolium Johanss., H. expallidiforme (Dahlst. ex. Stenstr.) Dahlst. var. oligasterum Johanss. & Sam. and H. insuccatum Johanss. var. occidentale Johanss. & Sam. respectively, when treated at the rank of species.     

中国高等植物多样性编目进展

中国高等植物多样性编目是“中国生物物种名录”项目中的重要组成部分, 旨在后植物志时代对中国高等植物的物种名录进行修订和年度更新。自2017年深圳国际植物学大会以来, 中国高等植物物种名录已进行了5次年度更新。名录参考最新的植物分类学研究进展, 对各大类群全面采用有分子证据的新分类系统, 收录5年来发表的中国植物新类群和新记录, 补充往年遗漏的部分类群, 并依据新的专科专属的修订结果对发生属间分类变动的类群进行了更新。目前的中国高等植物物种名录包含角苔类4科9属27种, 苔类62科170属1,081种94个种下等级, 藓类94科453属2,006种154个种下等级, 石松类3科12属165种4个种下等级, 蕨类38科177属2,215种228个种下等级, 裸子植物10科45属291种118个种下等级, 被子植物272科3,409属32,708种6,909个种下等级, 共计483科4,275属38,493种7,507个种下等级, 较5年前增加了19科270属2,334种。     

MOLECULAR SYSTEMATICS OF ECTOCARPUS AND KUCKUCKIA (ECTOCARPALES, PHAEOPHYCEAE) INFERRED FROM PHYLOGENETIC ANALYSIS OF NUCLEAR- AND PIASTTD-ENCODED DNA SEQUENCES

The phylogeny of Ectocarpus and Kuckuckia strains representing widely separated populations from both hemispheres was inferred from sequence analysis of the internal transcribed spacers of the nuclear ribosomal DNA (ITS 1—5.8S-ITS 2) and the spacer region in the plastid-encoded ribulose- bis -phosphate-carboxylase (RUBISCO) cistron (partial rbc L-spacer-partial rbc S ). Both sequences resulted in matching phylogenies, with the RUBISCO spacer region being most informative at the level of genera and species and the internal transcribed spacer sequences at the level of species and populations. Three major clades were formed by strains previously described by morphology and physiology as Kuckuckia, E. fasciculatus, and E. siliculosus, confirming the validity of these taxa . Ectocarpus and Kuckuckia are regarded as sibling taxa with respect to the outgroup species Feldmannia simplex, Hincksia mitchelliae, and Pilayella littoralis. The clade formed by sexual E. siliculosus strains and most asexual Ectocarpus strains was subdivided into several clades that are consistent with geographical races within E. siliculosus. The inferred phylogeny of Ectocarpus corresponds generally with results from cross-fertilization experiments, morphology, and lipid analysis. A hypothesis on the origin and dispersal of E. siliculosus suggests several natural dispersal events during periods of global cooling as well as recent and possibly anthropogenic dispersal events .