A new species of Carex section Ovales (Cyperaceae) occurring in the Ozark Mountain region

Carex ozarkana, a new species from Arkansas, Oklahoma, and extreme northeasternmost Texas is described and illustrated. It is a distinctive regional endemic of mineral soil wetlands most similar to the widespreadCarex albolutescens andC. longii but differing in its elongate, nodding inflorescences (on robust plants), prominently clavate spikes with elongate staminate bases, reddish brown pistillate and staminate scales, and wider achenes with a long apiculum.     

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.     

Carex tenuirostrata (Cyperaceae), a new species of section Confertiflorae from Zhejiang, China

Carex tenuirostrata, a new species of C. sect. Confertiflorae, is described from Pingyang and Putuo counties of Zhejiang province, China. It is morphologically similar to C. pseudodispalata, but differs in its androgynous lateral spikes; distended perigynia attenuately contracted into a short beak at the apex; ellipsoid-obovoid achenes contracted into a slender beak at the apex; and long-sheathed lower bracts. Scanning electron microscope (SEM) photographs of the achenes of the new species and C. pseudodispalata are provided for comparison.     

Genetic diversity and endemism in North American Carex section Ceratocystis (Cyperaceae)

Factors leading to endemism, and the evolutionary implications of endemism, can be explored by studying closely related taxa with variously restricted distributions. Such a model is provided by Carex section Ceratocystis (Cyperaceae); Carex cryptolepis, Carex sp. nov., and C. lutea are North American endemics with broad, restricted, and highly restricted distributions, respectively. The prediction that levels of genetic diversity are a consequence of distribution size was tested within a phylogenetic context using population level genetic variation at 18 allozyme loci. In contrast to expectations, mean proportion of loci polymorphic, number of alleles per polymorphic locus, and expected heterozygosity were significantly greater in C. lutea than either C. cryptolepis or Carex sp. nov. Although the possibility of a shift in breeding system, past introgression, or progenitor-derivative relationships could explain the relatively high levels of variation observed in C. lutea, these were dismissed on the basis of allozyme and nuclear ribosomal sequence data. We conclude that C. lutea maintains levels of genetic diversity typical of caespitose carices despite its narrow endemism and that the low levels of genetic variation in C. cryptolepis and Carex sp. nov. are likely the result of population fluctuations during Pleistocene glacial-interglacial cycles.     

Pollen morphology of selected Central European species from subgenera Vignea and Carex (Carex,Cyperaceae) and its relation to taxonomy

This study analysed the taxonomic utility of selected features of pollen grains of 45 Central European species of subgenera Vignea and Carex of the genus Carex. Seven quantitative features were analysed: [length of polar axis (P), length of equatorial axis (E), exine thickness on the pole (Exp), exine thickness on the equator plane (Exe), P/E ratio and the relative thickness of exine (Exp/P and Exe/E ratio)] and outline, shape and exine sculpture. Among the biometrical features, the most important for Carex taxonomy (particularly at the level of subgenus) is the thickness of exine (Exp). Marked differences in the morphological structure of inflorescences at the subgenus level are reflected in the differences of pollen features. Results of our studies clearly show an explicit (and not previously described in the palynological literature) taxonomic division of the genus Carex on the basis of pollen biometrical features into two units corresponding exactly to the subgenera (Vignea and Carex). However, the features of pollen grains analysed did not contribute any essential taxonomic criteria at levels lower than subgenus (e.g. sections). An exception was for C. disticha, where it was observed that the poroid areas occurred in pairs, a new feature for sedges. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 422–439.     

Environmental distribution of four Carex species (Cyperaceae) in an old-growth forest

We conducted an in-depth characterization of the range of micro-environments (1 m) in which four Carex species (C. backii, C. communis, C. plantaginea, and C. platyphylla) grow in the understory of an old-growth, deciduous forest in southern Québec, Canada. All four species occurred in significantly different micro-environments. Carex plantaginea was found at the wet end of a moisture gradient, in soils with high nitrate availability. Carex backii and C. platyphylla were found at the dry end of the moisture gradient, with C. backii occupying soils with higher phosphorus availability than C. platyphylla. Carex communis, the only ant-dispersed species studied, was found in the broadest range of environmental conditions. Our results suggest that environmental heterogeneity and interspecific microhabitat preferences are important for the maintenance of local species diversity in the forest understory, not only for common species as demonstrated in previous studies, but for infrequent species, and those within a functional group (upland Carex species). However, there was some evidence that the distributions of C. backii and C. communis were not in equilibrium with current environmental conditions, indicating that historical factors, such as dispersal and colonization events, may also have important effects on the distributions of these species and the maintenance of species diversity in old-growth forest.     

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。     

Two new species of Rhynchospora section Psilocarya (Cyperaceae)

Two new species ofRhynchospora sectionPsilocarya are described and illustrated.Rhynchospora eburnea is known from Amazonian Venezuela and nearby Brazil whileRhynchospora waspamensis is known only from eastern Zelaya, Nicaragua. These species are distinguished from most of the species of sectionPsilocarya by means of a key to the species found in the Americas south to and including the Amazon Basin.     

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.     

Allozyme variation and genetic relationships among species in the Carex willdenowii complex (Cyperaceae)

A taxonomic study by Naczi, Reznicek, and Ford (American Journal of Botany, 85, 434-447, 1998) has determined that three species (Carex willdenowii, C. basiantha, and C. superata) can be recognized within the C. willdenowii complex. To determine the amount of genetic divergence within and between these species, allozyme analyses were conducted on 14 populations distributed from Pennsylvania to eastern Texas. Seventeen loci were surveyed, 13 of which were polymorphic, with all populations being polymorphic at one or more loci. Interspecific genetic identities ranged from 0.560 (C. willdenowii and C. basiantha) to 0.807 (C. basiantha and C. superata). Alleles for the isozymes Aat-1, Dia-1, Idh-2, Mdh-2, Per-1, Pgm-1, and Pgm-2 served to distinguish C. willdenowii from C. basiantha and C. superata. Carex basiantha and C. superata were recognized by alleles of Mdh-2, Pgm-1, and Tpi-2. The genetic identities of populations within species were high and exceeded 0.957. A caespitose growth habit and perigynia in close proximity to the staminate flowers suggest adaptations for selfing and therefore low levels of heterozygosity. Paradoxically, the values for expected heterozygosities (Hexp) were always lower than those obtained by direct count (Hobs): F values were highly negative, indicating heterozygous excess. Disassortative mating and selection are discussed as possible mechanisms for maintaining heterozygous excess within populations.     

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.     

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.     

Species cohesion of an extremophyte (Carex angustisquama,Cyperaceae) in solfatara fields maintained under interspecific natural hybridization

Background and AimsHybridization is the main driver of plant diversification, and gene flow via hybridization has multifaceted effects on plant evolution. Carex angustisquama is an extremophyte that grows on soils heavily acidified by volcanism. Despite its habitat distinct from that of other species, this species is known to form interspecific hybrids, implying interspecific gene flow. It is crucial to verify the extent and direction of interspecific gene flow between C. angustisquama and closely related species to understand the evolutionary process of an extremophyte in solfatara fields.MethodsIn this study, expressed sequence tag–simple sequence repeat markers were utilized to infer the extent and direction of interspecific gene flow between C. angustisquama and closely related species.Key ResultsBayesian clustering and simulation analyses revealed that all individuals of the three hybrid species were classified into the first hybrid generation or first backcross to C. angustisquama; therefore, current interspecific gene flow is limited. Moreover, in the Bayesian inference of historical gene flow based on multispecies samples, the model that assumed no interspecific gene flow was the most strongly supported across all species pairs, including phylogenetically close but ecologically distinctive species pairs.ConclusionsOur results revealed that interspecific gene flow between C. angustisquama and its related species has been limited both currently and historically. Moreover, our results of Bayesian inference of historical gene flow indicated that extrinsic, rather than intrinsic, factors probably act as isolating barriers between Carex species, with hybrid breakdown via microhabitat segregation being the probable potential barrier. Overall, our findings provide insights into the evolutionary process of an extremophyte in solfatara fields and offer an important example of the mechanisms of diversification of the speciose genus Carex.