Significance of ecological vicariance and long-distance dispersal in the diversification of Carex sect. Spirostachyae (Cyperaceae)

Plant disjunctions have provided one of the most intriguing distribution patterns historically addressed by biogeographers. Carex sect. Spirostachyae (Cyperaceae) displays an interesting pattern of disjunction to evaluate these scenarios, with species occurring in the main continental landmasses and in oceanic islands of the two hemispheres. Internal transcribed spacer and 5'-trnK intron plastid gene sequences were analyzed to determine (1) the times of diversification using penalized likelihood, and (2) reconstructions of the regions using maximum likelihood and Bayesian approaches of origin of sect. Spirostachyae and internal main lineages. The times for the diversification of sect. Spirostachyae are dated to between the end of the Eocene and the Oligocene, whereas the two main lineages are dated to between the end of the Oligocene and the beginning of Miocene. The phylogenetic analyses reveal a Mediterranean-Eurasian center of differentiation for sect. Spirostachyae and subsection Spirostachyae, whereas no clear, single ancestral area could be inferred for subsection Elatae. Both long-distance dispersal and ecological vicariance appear to have been involved in the evolutionary history of the disjunct distribution of the main lineages of sect. Spirostachyae. These organisms appear to have a special ability to colonize remote areas (through transoceanic and interhemispherical colonizations), but special long-distance dispersal mechanisms are not evident.     

Growth pattern of Carex arenaria (Cyperaceae)

The origin of the orthotropic aerial shoot from the plagiotropic, horizontal rhizome of Carex arenaria L. (Cyperaceae) was studied. A previously proposed adnation or fusion of the first internode of the orthotropic shoot and the fourth internode of the plagiotropic shoot could not be anatomically confirmed. The rhizome branches sympodially and the point of branching results from intercalary meristematic activity in the plagiotropic shoot.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 145–151.     

福建薹草(莎草科)的考证

在整理和鉴定浙闽薹草属Carex植物的过程中,发现福建薹草Cares fokienensis Dunn较早发表但并未被注意。通过文献查阅、模式标本研究和野外观察,认为闽清薹草C.minqingensis Z．P.Wang、九仙山薹草C jiuxiangshanensis L.K．Dai＆Y.Z．Huang、苍绿薹草C.pallideviridis K．L．Chv（裸名）在果囊和雌花鳞片等鉴别性特征上与福建薹草并没有区别,应该处理为福建薹草的异名。     

Systematics of the Carex aquatilis and C. lenticularis lineages: Geographically and ecologically divergent sister clades of Carex section Phacocystis (Cyperaceae)

Carex aquatilis is a highly diverse and geographically widespread member of one of the largest genera of flowering plants, Carex, and is ideally suited for the study of the role of hybridization and niche partitioning in ecological speciation. Phylogenetic analyses of nuclear ITS and ETS 1f and chloroplast psbA-trnH DNA sequences support the monophyly of a broadly defined Carex aquatilis-Carex lenticularis lineage, which includes C. aquatilis and C. lenticularis and their allies within section Phacocystis. However, neither taxon is monophyletic as currently circumscribed. The C. aquatilis lineage includes C. aquatilis and four morphologically and molecularly distinct salt-tolerant maritime taxa with which C. aquatilis s.s. is reported to form stabilized homoploid hybrids. The C. lenticularis lineage includes a paraphyletic C. lenticularis and seven allied species from both the New and Old World. The data provided here allow recognition of four species within the North American endemic C. lenticularis and suggest a neotropical origin for the C. lenticularis lineage with subsequent radiation and divergence through northwestern North America to Asia and via northeastern North America to Europe and southern South America. Evolutionary rate analyses indicate an origin for the C. aquatilis-C. lenticularis group around 1.89 million years ago during the early Pleistocene.     

Mycorrhizal status of the genus Carex (Cyperaceae)

The Cyperaceae have generally been considered nonmycorrhizal, although recent evidence suggests that mycotrophy may be considerably more widespread among sedges than was previously realized. This study surveyed 23 species of Carex occurring in upland and wetland habitats in northeastern Illinois. Mycorrhizal infection by arbuscular fungi was found in the roots of 16 species of Carex and appears to occur in response to many factors, both environmental and phylogenetic. While some species appear to be obligately nonmycorrhizal, edaphic influences may be responsible for infection in others. In five of the seven Carex species that were nonmycorrhizal, a novel root character, the presence of bulbous-based root hairs, was identified. The taxonomically patchy distribution of the distinctive root hair trait suggests that these structures may have evolved several times within the genus. Evidence of multiple independent origins of the root hair trait lends support to the hypothesis that root hairs represent an adaptation to nonmycotrophy. Although taxonomic position does seem to be of importance in determining the mycorrhizal dependence of sedges, the pattern may be a patchwork of both mycorrhizal clades and clades that have adapted to the nonmycorrhizal state.     

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.     

Geographical vs. ecological diversification in Carex section Phacocystis (Cyperaceae): Patterns hidden behind a twisted taxonomy

Carex section Phacocystis (Cyperaceae) is one of the most diverse and taxonomically complex groups of sedges (between 116 and 147 species), with a worldwide distribution in a wide array of biomes. It has a very complicated taxonomic history, with numerous disagreements among different treatments. We studied the biogeography and niche evolution in a phylogenetic framework to unveil the relative contribution of geographical and ecological drivers to diversification of the group. We used a large species sampling of the section (82% of extant species) to build a phylogeny based on four DNA regions, constrained with a phylogenomic HybSeq tree and dated with six fossil calibrations. Our phylogenetic results recovered section Phacocystis s.s. (core Phacocystis) as sister to section Praelongae. Ancestral area reconstruction points toward the N Pacific as the cradle for the crown diversification of section Phacocystis during the Middle Miocene. Wide distributions were recurrently inferred across deep nodes. Large Northern Hemisphere lineages with geographical congruence were retrieved, pointing toward the importance of allopatric divergence at deep phylogenetic levels, whereas within-area speciation emerges as the predominant pattern at shallow phylogenetic level. The Southern Hemisphere (Neotropics, SW Pacific) was colonized several times from the Northern Hemisphere. The global expansion of Carex section Phacocystis did not entail major ecological changes along the inner branches of the phylogeny. Nevertheless, ecological differentiation seems to gain importance toward recent times.     

A tale of worldwide success: Behind the scenes of Carex (Cyperaceae) biogeography and diversification

The megadiverse genus Carex (c. 2000 species, Cyperaceae) has a nearly cosmopolitan distribution, displaying an inverted latitudinal richness gradient with higher species diversity in cold‐temperate areas of the Northern Hemisphere. Despite great expansion in our knowledge of the phylogenetic history of the genus and many molecular studies focusing on the biogeography of particular groups during the last few decades, a global analysis of Carex biogeography and diversification is still lacking. For this purpose, we built the hitherto most comprehensive Carex‐dated phylogeny based on three markers (ETS–ITS–matK), using a previous phylogenomic Hyb‐Seq framework, and a sampling of two‐thirds of its species and all recognized sections. Ancestral area reconstruction, biogeographic stochastic mapping, and diversification rate analyses were conducted to elucidate macroevolutionary biogeographic and diversification patterns. Our results reveal that Carex originated in the late Eocene in E Asia, where it probably remained until the synchronous diversification of its main subgeneric lineages during the late Oligocene. E Asia is supported as the cradle of Carex diversification, as well as a “museum” of extant species diversity. Subsequent “out‐of‐Asia” colonization patterns feature multiple asymmetric dispersals clustered toward present times among the Northern Hemisphere regions, with major regions acting both as source and sink (especially Asia and North America), as well as several independent colonization events of the Southern Hemisphere. We detected 13 notable diversification rate shifts during the last 10 My, including remarkable radiations in North America and New Zealand, which occurred concurrently with the late Neogene global cooling, which suggests that diversification involved the colonization of new areas and expansion into novel areas of niche space.     

Carex, subgenus Carex (Cyperaceae) – A phylogenetic approach using ITS sequences

To evaluate the sectional classification in Carex, subgenus Carex, the ITS region of 117 species belonging to 32 sections was analyzed with Neighbor Joining (NJ) and Markov chain Monte Carlo (MCMC) methods. In our analyses (1) species of subgenus Indocarex appear as a statistically well supported group within subgenus Carex. (2) The representatives of sections Vesicariae, Hirtae, Pseudocypereae, Ceratocystis, Spirostachyae, Bicolores, Paniceae, Trachychlaenae, Scirpinae, Atratae and Albae group in statistically supported clades with higher support in MCMC than in NJ. (3) C. rariflora clusters with representatives of section Limosae, however only weakly supported. (4) Taxa of section Phacocystis are divided in two statistically supported subclusters that are closely related to a core group of section Hymenochlaenae. (5) Species of sections Montanae, Pachystylae, Digitatae, Phacocystis, Rhomboidales, Careyanae and Frigidae are segregated into two or more clusters each. (6) Five species of section Frigidae cluster together, whereas the seven others are in scattered positions. Based on these results, delimitation of sections is discussed.     

Allozyme differentiation between the lowland Carex capitata and the alpine Carex arctogena (Cyperaceae) in Scandinavia

Within Scandinavia, Carex capitata s.l, is usually divided into two taxa: the lowland to lowalpine C. capitata s.s., and the alpine C. arctogena. The systematic status of these taxa is uncertain, reflected by their taxonomic treatment in different floras. In this study, allozymes were used to assess the degree of genetic differentiation between and within taxa. Ten populations of Carex capitata s.s. and eight of C. arctogena, from Norway and Sweden, were included in the study. Sixteen loci in eleven different enzyme systems were analysed: five were variable within or between taxa, three could be interpreted in all plants. In all, 20 different alleles were found. Four of these alleles were confined to C. capitata s.s., and two to C. arctogena. In C. capitata s.s., three loci (18%) were polymorphic, while all loci were monomorphic in C. arctogena. The number of alleles per polymorphic locus was 2.2 in C. capitata s.l. The taxa had completely different alleles at two loci {Pgi-2, Pgm-1). Nei's genetic identity was 0.86 between taxa, and ranged between 0.95 and 1 in pairwise comparisons of populations of Carex capitata s.s., while it was 1 in all comparisons between populations of C. arctogena. The distinction of C. capitata s.s. and C. arctogena as separate species agrees well with the amount of differentiation found between other congeneric species. It is suggested that the low genetic variation observed in both taxa is due to ancient founder events. The difference in genetic variability between taxa may be due to differences in immigration history: whereas C. capitata s.s. may have colonized Scandinavia from several close refugia after the last glaciation, C. arctogena may have reached Scandinavia after long-distance dispersal of a limited number of propagules.     

Sectional limits and phylogenetic considerations in Carex section Ceratocystis (Cyperaceae)

Carex L. sectionCeratocystis Dumort. is distinct from the closely related sectionSpirostachyae (Drejer) L. Bailey. Differences in pistillate spike morphology, achene shape, and perigynium epidermis are augmented by chemical, cytological, and micromorphological evidence. Character compatibility analysis has been used to reconstruct the phylogenetic history of the sedges in sectionCeratocystis. The resulting network is shown to be robust when tested with cytological, ecological, and micromorphological data not included in its construction. Agmatoploidy has been the dominant process of chromosomal evolution in the section.     

莎草科薹草属黑穗薹草组的植物地理学讨论

根据黑穗薹草组植物的外部形态、细胞染色体特征与地理分布之间的关系,结合地质历史事件推测了黑穗薹草组Sect.Racemosae G.Don.的起源、进化和迁移等问题。推测黑穗薹草组可能在早第三纪的始新世起源于我国的喜马拉雅-横断山地区;可能的迁移路径为：由起源中心向北部迁移至欧洲、中亚、俄罗斯西伯利亚地区,并通过格陵兰岛和白令陆桥到达北美洲地区;经过第四纪冰川后,一些分布于北美洲的物种又通过白令陆桥回迁到亚洲东部的俄罗斯远东地区,最终形成该组植物的现代分布格局。     

Taxonomy of the Carex flava complex (Cyperaceae) in Finland

The morphological variation in the Carex flava complex (sect. Ceratocystis ) in Finland, particularly taxa described by Alvar Palmgren as endemic to northern Europe, was studied using numerical analyses. Discriminant analyses were used to maximize the separation of the taxa. However, the separation of the taxa described by Palmgren was weak. The taxonomic treatment of the C. flava complex in Finland based on morphology, ecology, distribution and hybridization of the taxa is presented. Four species are recognized: Carex flava (the lectotype is selected here), C. lepidocarpa, C. demissa and C. viridula. C. lepidocarpa includes ssp. lepidocarpa and ssp. jemtlandica . In the polymorphic C. viridula , three taxa are recognized: var. viridula , var. bergrothii and var. pulchella . A key to the taxa is presented. The distribution of the taxa in Finland is presented, and their ecology is briefly discussed.     

Chromosome counts in the Carex flava complex (Cyperaceae) in Finland

Chromosome numbers from 17 Finnish populations of six taxa in the Carex flava complex were examined. Special attention was given to taxa claimed to be endemic to Fennoscandia. The meiotic counts for C. flava n = 30, C. viridula var. viridula n = 35 and var. pulchella n = 35 are in accordance with previous reports. The counts for C. jemtlandica n = 34, C. bergrothii n = 35, and C. kotilaini n = 34 are new. The hybrid C. flava × kotilaini reveals severely disturbed meiosis with many uni- and multivalents. The results are briefly discussed in relation to morphological data. Carex jemtlandica should evidently be included in C. lepidocarpa. Carex bergrothii and C. kotilaini belong to the polymorphic C. viridula , but the latter probably consitute a specific cytotype.     

Nonuniform processes of chromosome evolution in sedges (Carex: Cyperaceae)

Holocentric chromosomes-chromosomes that lack localized centromeres-occur in numerous unrelated clades of insects, flatworms, and angiosperms. Chromosome number changes in such organisms often result from fission and fusion events rather than polyploidy. In this study, I test the hypothesis that chromosome number evolves according to a uniform process in Carex section Ovales (Cyperaceae), the largest New World section of an angiosperm genus renowned for its chromosomal variability and species richness. I evaluate alternative models of chromosome evolution that allow for shifts in both stochastic and deterministic evolutionary processes and that quantify the rate of evolution and heritability/phylogenetic dependence of chromosome number. Estimates of Ornstein-Uhlenbeck model parameters and tree-scaling parameters in a generalized least squares framework demonstrate that (1) chromosome numbers evolve rapidly toward clade-specific stationary distributions that cannot be explained by constant variance (Brownian motion) evolutionary models, (2) chromosome evolution in the section is rapid and exhibits little phylogenetic inertia, and (3) explaining the phylogenetic pattern of chromosome numbers in the section entails inferring a shift in evolutionary dynamics at the root of a derived clade. The finding that chromosome evolution is not a uniform process in sedges provides a novel example of karyotypic orthoselection in an organism with holocentric chromosomes.     

Macroevolutionary insights into sedges (Carex: Cyperaceae): The effects of rapid chromosome number evolution on lineage diversification

Changes in holocentric chromosome number due to fission and fusion have direct and immediate effects on genome structure and recombination rates. These, in turn, may influence ecology and evolutionary trajectories profoundly. Sedges of the genus Carex (Cyperaceae) comprise ca. 2000 species with holocentric chromosomes. The genus exhibits a phenomenal range in the chromosome number (2n = 10 − 132) with almost not polyploidy. In this study, we integrated the most comprehensive cytogenetic and phylogenetic data for sedges with associated climatic and morphological data to investigate the hypothesis that high recombination rates are selected when evolutionary innovation is required, using chromosome number evolution as a proxy for recombination rate. We evaluated Ornstein–Uhlenbeck models to infer shifts in chromosome number equilibrium and selective regime. We also tested the relationship between chromosome number and diversification rates. Our analyses demonstrate significant correlations between morphology and climatic niche and chromosome number in Carex. Nevertheless, the amount of chromosomal variation that we are able to explain is very small. We recognized a large number of shifts in mean chromosome number, but a significantly lower number in climatic niche and morphology. We also detected a peak in diversification rates near intermediate recombination rates. In combination, these analyses point toward the importance of chromosome evolution to the evolutionary history of Carex. Our work suggests that the effect of chromosome evolution on recombination rates, not just on reproductive isolation, may be central to the evolutionary history of sedges.     

浙江薹草属（莎草科）新分类群

摘要描述及图解了浙江薹草属Carex 5新种和1新变种,它们是：灰帽薹草组sect. Mitratae Kükenth.的天目山薹草Carex tianmushanica C. Z. Zheng & X. F. Jin和拟三穗薹草C. pseudotristachya X. F. Jin & C. Z. Zheng;匏囊薹草组sect. Lageniformes (Ohwi) Nelmes的密毛薹草C. densipilosa C. Z. Zheng & X. F. Jin;菱形果薹草组sect. Rhomboidales Kükenth.的浙南薹草C. austrozhejiangensis C. Z. Zheng & X. F. Jin和朝芳薹草C. chaofangii C. Z. Zheng & X. F. Jin;疏花薹草组sect. Laxiflorae Kunth的无芒长嘴薹草C. longerostrata C. A. Mey. var. exaristata X. F. Jin & C. Z. Zheng。     

Clonal structure and hybrid susceptibility to a smut pathogen in microscale hybrid zones of northern wetland Carex (Cyperaceae)

Interspecific hybrid taxa, especially those with the potential for clonal spread, may play important roles in community dynamics and plant-pathogen interactions. This study combines the mapping of clonal structure for two rhizomatous sedges (Carex limosa, C. rariflora) and their nearly sterile interspecific hybrid with an investigation of the relationship between these taxa and a nonsystemic floral smut pathogen (Anthracoidea limosa) in six subarctic fens in Nouveau-Québec, Canada. We used allozyme polymorphisms in 14 of 18 putative loci to confirm hybrid identification and to distinguish among genotypes for mapping. The incidence of A. limosa was 5-20 times greater on hybrids than on parental taxa across all sites at two spatial scales (intensive extent = 10.5 m(2), extensive extent = entire fens). Spatial autocorrelation was detected in smut incidence; however, its statistical removal did not alter the strong association between hybrids and smut infection. Smut incidence on both C. limosa and hybrids was greater when they were growing in areas of high hybrid density. Our study provides evidence that disease can help maintain boundaries between species. We suggest explanations for hybrid susceptibility and provide evidence for a model in which hybrids act as a source for reinfection for all three taxa during subsequent years.     

Development of aerenchyma in roots and rhizomes of Carex rostrata (Cyperaceae)

Aerenchyma development in Carex rostrata was studied using light and scanning electron microscopy. Specimens were collected at two locations in southern Finland. Examination showed the beginning of aerenchyma development in the cortex of roots at the distance of 30–45 mm from the apex and it was fully developed at 75–90 mm from the apex. First, schizogenous cavities were formed in the cortex, and the sequence continued with a collapse of tangential cell walls in the cortex, leaving radial strands of cells intact and leading to a structure resembling a cobweb in cross section. In the rhizomes some of the radial cell walls disintegrated in the cortex, resulting in radial rows of cells without any tangential connections. No diaphragms were observed in the rhizome. There were no direct contacts between the gas spaces of the roots and the rhizomes. The significance of aerenchyma for metabolic processes is discussed.