Patterns of allozyme and morphological differentiation in the Carex flava complex (Cyperaceae) in Fennoscandiaxs

Allozyme and morphological differentiation patterns within the Carex flava complex in Fennoscandia were described by means of multivariate analyses. As a result of these studies, and considering also data on ecology, distribution and hybridization, the following taxa were recognized: C. flava with no infraspecific taxa, C. lepidocarpa with two subspecies, ssp. lepidocarpa and ssp. jemtlandica, C. demissa with no infraspecific taxa, and C. oederi with three varieties, var. oederi , var. bergrothii , and var. pulchella. Carex lepidocarpa ssp. jemtlandica was allozymically divergent from ssp. lepidocarpa , indicating a long period of isolation of the two subspecies. A poor morphological separation in some localities is apparently due to secondary contacts in connection with recolonization of Fennoscandia after the last ice age, resulting in the formation of fertile hybrids and back-crosses. The subdivision of C. oederi into three varieties is supported by ecological data and it is suggested that they have originated by parallel ecotype formation under similar selective regimes in different areas. However, the varietal limits are not supported by differentiation in neutral (allozyme) characters. A phylogenetic hypothesis was constructed by means of neighbour-joining based allozyme data and it was indicated that C. demissa may be more closely related to C. lepidocarpa than to C. oederi. Previous treatments that have reduced the C. flava complex to one or two species may have been based on inappropriate species concepts, too coarse methods of analysis of morphological data, ignorance of patterns of co-occurrence and hybridization, or a combination of these factors.     

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.     

Within-species variation among populations of the Carex flava complex as a function of habitat conditions

Background: Variable habitat conditions contribute to morphological variability that plays a substantial part in evolution of plants. Understanding the extent of phenotypic plasticity has important implication for assigning individuals to taxa.

Aims: I analysed morphological variability among populations within species of the Carex flava group and tested to what extent morphological variability was associated with habitat conditions.

Methods: A total of 571 specimens from 20 populations of four species from the C. flava complex were sampled in Poland and tested by Discriminant Function Analysis (DFA). The relationship between morphological characters and habitat factors was explored by means of the redundancy analysis (RDA).

Results: Variability of the generative and vegetative traits was fairly similar in Carex lepidocarpa, C. flava and Carex demissa, while it was somewhat higher in Carex viridula. The morphological traits were mostly related to soil organic matter content, calcium and carbonate, as well as to habitat fertility (C:N ratio), elevation and habitat disturbance. The results obtained do not support the separation of C. viridula var. pulchella from C. viridula var. viridula.

Conclusions: Phenotypic variability in the species of the C. flava complex is related to habitat conditions and this can lead to the differentiation of morphotypes within species.     

Studies in African Cyperaceae 27. Miscellaneous new taxa and combinations

One new form of Fuirena Rottb., one new subspecies of Schoenoplectus Palla and one new variety of Carex L. are described from East Africa, viz. Fuirena leptostachya Oliv. f. nudiflora Lye, Schoenoplectus lateriflorus (Gmel.) Lye ssp. laevinux Lye and Carex bequaertii De Wild. var. maxima Lye. In addition, the following new combinations are made: Fuirena ciliaris (L.) Roxb. f. apetala (Wingfield) Lye, F. stricta Steudel ssp. chlorocarpa (Ridley) Lye, Schoenoplectus rhodesicus (Podlech) Lye, 5. corymbosus (Roem. & Schult.) J. Rayn. var. brachyceras (A. Rich.) Lye, S. confusus (N.E. Br.) Lye var. rogersii (N.E. Br.) Lye, S. erectus (Poiret) J. Rayn, ssp. raynalii (Schuyler) Lye, S. erectus (Poiret) J. Rayn. ssp. sinuatus (Schuyler) Lye, Eleocharis retroflexa (Poiret) Urban ssp. subtilissima (Nelmes) Lye, Machaerina Vahl subgen. Baumea (Gaud.) Lye, Machaerina flexuosa (Bock.) Kern ssp. laevinux (J. Rayn.) Lye, M. montana (J. Rayn.) Lye, M. flexuosa (Böck.) Kern ssp. polyanthemum (Kük.) Lye, Scleria lagoënsis Böck. ssp. canaliculato–triquetra (Böck.) Lye, 5. distans Poiret var. glomerulata (Oliv.) Lye, S. distans Poiret var. chondrocarpa (Nelmes) Lye, S. melanotricha A. Rich. var. grata (Nelmes) Lye, Carex echinochloë Kunze ssp. nyasensis (C. B. Clarke) Lye, C. conferta A. Rich. var. lycurus (K. Schum.) Lye, C. cognata Kunth var. abyssinica (Chiov.) Lye and C. cognata Kunth var. congolensis (Turrill) Lye.     

Allozyme variation and racial differentiation in Swedish Carex lepidocarpa s.l. (Cyperaceae)

Two morphological races have previously been recognized within the sedge, Carex lepidocarpa , in Sweden. These largely allopatric races are accorded specific status, as C. lepidocarpa s.s. and C. jemtlandica , in Scandinavian floras. A study of allozyme variation in populations from 80 Swedish sites supports the morphological evidence for racial differentiation within C. lepidocarpa. The two races differ from each other in terms of allele frequencies at polymorphic loci and also show different levels of within-population genetic diversity. Material that is morphologically referable to C. lepidocarpa s.s. is characterized by relatively high levels of allozyme variation, both within and between populations. Carex lepidocarpa s.s. is widespread in southern Sweden. In contrast, material that is morphologically assignable to C. jemtlandica shows low levels of within-population genetic diversity, and there is little differentiation between the geographically separated isolates of C. jemtlandica in northern Sweden and on the Baltic island of Gotland. The high degree of morphological similarity and moderate levels of genetic differentiation between the two races within C. lepidocarpa indicate that it is more appropriate to recognize the races as subspecies than as species. The low levels of genetic variation in C. jemtlandica , both within and between populations, suggest that C. jemtlandica may have arisen from C. lepidocarpa (or a near ancestor of C. lepidocarpa) as a result of population fragmentation and isolation in glacial refugia, or during the process of post-glacial colonization of Scandinavia. Lack of allozyme evidence for extensive hybridization between the two races of C. lepidocarpa , despite their ability to hybridize freely where their ranges overlap at present, supports the suggestion that the two races have had separate post-glacial histories.     

中国薹草属(莎草科)研究随记(Ⅳ).隐匿薹草组的分类修订(英文)

隐匿薹草组(Carex sect.Infossae)为最近建立的薹草属(Carex)的组,当时报导分布于我国安徽和江苏,有2种和1变种,即隐匿薹草(C.infossa)、显穗薹草(变种)(C.infossa var.extensa)和矮秆薹草(C.minuticulmis)。根据描述,另有15种和2变种应归属本组,这些分类群具有较为一致的特征:果囊椭圆球状卵球形或卵球形,通常疏被短毛或近无毛,先端渐狭成中等长的喙,喙口具2小齿;瘦果卵球形,先端钝或微凹。通过文献考证、标本查阅、野外采集,并结合果囊和瘦果的扫描电镜观察,对隐匿薹草组所涉及的种(含种下类群)进行了分类修订。该组仅有2种:百里薹草(C.blinii)和尖叶薹草(C.oxyphylla)。将Carex elmeri、C.granifera、矮秆薹草(C.minuticulmis)、上海薹草(C.shanghaiensis C.blinii subsp.shanghaiensis)、C.sublateralis、C.taihokuensis和锐果薹草(C.tatsutakensis)并入百里薹草;将沟囊薹草(C.canaliculata)、樟木薹草(C.changmuensis)、C.distantiflora、隐匿薹草(C.infossa)、显穗薹草(C.infossa var.extensa)、C.lateralis、C.loheri、无芒长嘴薹草(C.longerostrata var.exaristata)、城湾薹草(C.longerostrata var.hoi)、C.lyi、和平菱果薹草(C.macrandrolepis)和C.sharyotensis并入尖叶薹草。     

Morphological differentiation among closely related species with disjunct distributions: a case study of Mediterranean Cyclamen L. subgen. Psilanthum Schwarz (Primulaceae)

In Cyclamen subgenus Psilanthum , previous authors have recognized three species ( C. repandum, C. balearicum and C. creticum ), with C. repandum subdivided into three subspecies (ssp. repandum , ssp. peloponnesiacum and ssp. rhodense ), and two varieties (var. vividum and var. peloponnesiacum ) within C. repandum ssp. peloponnesiacum . To examine the validity of this classification and the degree of morphological differentiation among closely related endemic taxa with disjunct distributions, we undertook analyses of three quantitative and eight qualitative morphological traits of flowers and leaves, most of which have previously been used without statistical comparison in the classification of the different taxa. Cyclamen balearicum showed high levels of morphological differentiation from all other taxa in subgenus Psilanthum consistent with its specific status. In contrast, morphological differentiation between C. creticum and subspecies of C. repandum was similar to that among the three subspecies of C. repandum . When compared with a recent molecular phylogeny of this subgenus, our results suggest that C. creticum may best be described as a geographical subspecies of C. repandum . The two varieties of C. repandum ssp. peloponnesiacum showed few significant differences for individual traits and their overall morphology is very similar to each other. The morphology of plants at three sites on Corsica strongly suggest, consistent with molecular data, the occurrence of hybrid populations between C. balearicum (outside of its previously recognized distribution) and local C. repandum .  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 133–144.     

A subspecific taxonomic study on Asplenium trichomanes L. from China

With the evidence from palynological, cytological, ecological and morphological data, the subspecific taxonomy of Asplenium trichomanes L. from China is carried out. Four subspecies and one variety, i.e.A. trichomanes L. ssp. trichomanes, A. trichomanes L. ssp. inexpectans Lovis, A. trichomanes L. ssp. quadrivalens D. E. Meyer emend. Lovis, A. trichomanes L. ssp. pachyrachis (Christ) Lovis et Reichst. and A. trichomanes L. var. harovii Moore emend. Midle are recognized from China. The distribution of each subspecies and variety is also presented. Some fragments of a type specimen named as A. trichomanes L. var. centrochinense Christ in PE are found to be different from the known taxa of A. trichomanes L. complex both morphologically andcytologically, and therefore are regarded representing a new species.     

Shoot and leaf demography of Carex curvula ssp. Curvula and Carex curvula ssp. rosae in the Central Alps

Abstract. Populations of Carex curvula ssp. curvula, C. curvula ssp. rosae and their intermediate form were investigated in the Central Alps over a three-year period. The closely related taxa showed a different dominance behaviour in their respective communities. This may be caused by different growth strategies and a different reproduction biology. Therefore, the main aim of the study was to compare the demography of the three taxa. Shoot and leaf turnover, flower and seed production and population half-lives were determined. Differences in growth dynamics were less pronounced between the two species of Carex curvula than between these taxa and their intermediate form which showed the highest shoot turnover, highest number of fertile inflorescences and highest number of seeds per inflorescence. Carex curvula ssp. rosae showed a slightly higher shoot and leaf production but a lower reproduction capacity than Carex curvula ssp. curvula. Recruitment of populations of the three Carex-forms was only by vegetative shoots. Flowering had a striking effect on the shoot dynamics of the species grown in the grassland sites: up to about 70 % of all dead shoots could be identified as dead flowering shoots from the previous year. The different dominance behaviour of the three taxa could not be explained by their demographical features. Interspecific interactions and the occurrence of microniches might affect the growth and reproduction processes of the taxa.     

Morphology and taxonomy of the genus Cochlearia (Brassicaceae) in Northern Scandinavia

The morphological variation between populations of Cochlearia in Northern Scandinavia has been analysed. The variation is correlated to ecology, and four ecotypes are recognized. Based on the morphological differentiation of the ecotypes and the results of crossing experiments, the populations are referred to the following three taxa: C. officinalis ssp. officinalis which includes the common beach ecotype and the bird cliff ecotype; C. officinalis ssp. integrifolia stat. nov. which includes the spring ecotype growing in inland localities up to about 600 m a.s.l. in cold more or less eutrophic springs, and C. officinalis ssp. norvegica ssp. nov. which includes the estuary ecotype, growing in poor, often inundated habitats near river outlets at the fjord ends. All the differentiation has taken place at the tetraploid level (2n = 24). The subspecies are kept more or less distinct due to ecological and seasonal isolating mechanisms rather than internal, reproductive barriers. A key to the subspecies and distribution maps for ssp. integrifolia and ssp. norvegica are presented.     

Studies in African Cyperaceae 21. New taxa and combinations in Kyllinga Rottb.

Eight new species, one new subspecies and one new variety of Kyllinga are described from tropical Africa, viz. Kyllinga brunneoalba Lye, K. afropumila Lye, K, albogracilis Lye, K. aureovillosa Lye, K. alba–purpurea Lye, K. rhizomafragilis Lye, K. microbracleata Lye, K. afro–occidentalis Lye, K. erecta Schumacher ssp. albescens Lye, and K. melanosperma Nees var. hexalata Lye. In addition nine new combinations are made, viz. Kyllinga peteri (Kükenthal) Lye, K. bigibbosa (Fosberg) Lye, K. odorata Vahl ssp. appendiculata (K. Schum.) Lye, K. alba Nees ssp. ascolepidioides (H. Cherm.) Lye, K. nervosa Steudel ssp.flava (C.B.C1.) Lye, K. melanosperma Nees ssp. elata (Steudel) Lye, K. brevifolia Rottb. ssp. lurida (Kükenthal) Lye, K. bre–vifolia Rottb. ssp. intricata (H. Cherm.) Lye, and K. comosipes (Mattf. & Kükenthal) Agnew & Hanid ssp. decolorans (Kükenthal) Lye.     

Allozyme variation in Japanese species ofChionographis (Liliaceae)

Allozyme variation was examined in three diploid taxaChionographis japonica var.japonica, var.kurokamiana, andC. koidzumiana and three tetraploid taxaC. japonica var.kurohimensis, ssp.hisauchiana, and ssp.minoensis. Results show thatC. japonica var.kurokamiana is genetically closer toC. koidzumiana than to var.japonica. In the tetraploid taxa, fixed heterozygosities were found at several loci, and this supports the hypothesis that these taxa are allotetraploids. Furthermore, the tetraploid taxa have many unique alleles not found in the diploid taxa. This suggests that sufficient time has passed since the origin of tetraploids for new mutations to have been fixed.     

The acaulescent and succulent species of Dorstenia sect. Kosaria (Moraceae) from NE tropical Africa and Arabia

The acaulescent and succulent species of Dorstenia sect. Kosaria in NE tropical Africa include the well defined D. ellenbeckiana and the two polymorphic taxa, the D. barnimiana complex and the D. foetida complex. The D. barnimiana complex is treated as two species, D. barnimiana Schweinf. and D. tropaeolifolia (Schweinf.) Bureau. One specific name and four varietal names are reduced to synonymy. The D. foetida complex is treated as one species, D. foetida (Forssk.) Schweinf., with two subspecies, ssp. foetida and ssp. lancifolia (Rendle) Friis comb, nov., of which the former is further divided into var. foetida and var. obovata (A. Rich.) Schweinf. & Engl. Three species arc reduced to synonymy of D. foetida. The distribution of all the species is mapped.     

Taxonomy and geographic distribution of <Emphasis Type="Italic">Carex lucorum</Emphasis> var. <Emphasis Type="Italic">austrolucorum</Emphasis> (section <Emphasis Type="Italic">Acrocystis</Emphasis>, Cyperaceae)

Carex lucorum currently comprises two morphologically similar varieties. We revise the taxonomy of C. lucorum, and map the distribution of the taxa. Based on multivariate analyses and analyses of variance of measured characters, allopatry, and previous studies revealing differences in achene micromorphology, chromosome differences, and flavonoid chemistry, we elevate C. lucorum var. austrolucorum to species status as C. austrolucorum. We provide an identification key to C. lucorum and its closest relatives.     

A taxonomìc revision of Hordeum sect. Critesion

Eight taxa are treated, viz. Hordeum pubiflorum ssp. pubiflorum (2n=14), ssp. halophilum (comb. nov.) (2n=14), H. comosum (2n=14), H. jubatum ssp. jubatum (2n = 28), H. jubatum ssp. intermedium (2n = 28), H. lechleri (2n=42), H. procerum (2n=42), and H. arizonicum (2n=42). Information about morphological variation, cytology, and genome relationship, ecology and distribution is presented.     

A revision of the genus Lasianthus (Rubiaceae) in Africa, excluding Madagascar

23 species, one subspecies and three varieties have been described or named for the pan-tropical genus Lasianthus Jack (Rubiaceae) in continental Africa. In the present revision thirteen species, five subspecies and one variety are recognized; additionally two hybrids are proposed. Of the previously named taxa, the subspecies is raised to species rank whereas two species are degraded to subspecies and variety status, respectively. Five species are synonymised including two varieties, and four are left with uncertain status, including one variety. The distribution of the genus in Africa consists of two distinct elements: one component is found in the Guineo-Congolian regional centre of endemism and one in the eastern part of Afiomontane Archipelago-like regional centre of endemism. A centre of species diversity is found in Eastern Arc Mts., where six endemic species occur together with L. laxinervis and the more widespread L. kilimandscharicus. Four of the species are strict endemics of the Uluguru Mts. In addition to the presentation of the new section: Lasianthus Sectio Membranacei, sect. nov., this revision contain two hybrids from Uluguru Mts.: L. macrocalyx × pedunculatus and L. cereifizorus × pedunculatus, and following new taxa and new combinations: L. africanus ssp. biokoensis, subsp. nov.; L. africanus ssp. mayumbensis, comb. et stat. nov.; L. kilimandscharicus ssp. glabrescens, ssp. nov.; L. kilimandscharicus ssp. glabrescens var. xanthospermus, comb. et stat. nov.; L. kilimandscharicus ssp. hirsutus, ssp. nov.; L. laxinervis, stat. nov.; L. pedunculatus ssp. angustisepalus, ssp. nov.     

Synopsis of Turkish Acantholimon Boiss. (Plumbaginaceae)

A synopsis of Acantholimon in Turkey is given on the basis of a revisional study conducted in Turkey by the authors. An account of 52 species, ten subspecies, and 17 varieties is given, including keys to the species, subspecies, and varieties, and a general indication of their distribution, as well as their conservation status in Turkey. Seven new taxa, namely A .  ekimii sp. nov. , A. capitatum ssp. sivasicum ssp. nov. , A. calvertii var. glabrum var. nov. , A. huetii var. breviscapum var. nov. , A. acerosum ssp. longibracteolorum ssp. nov. , A. lycaonicum ssp. cappadocicum ssp. nov. , and A. halophilum var. coloratum var. nov. , are described, together with a few new combinations. A. wiedemannii , A. iconicum , and A. lycaonicum , described from Turkey in the past, but placed under other species as synonyms in the Flora of Turkey , are recognized once again as good species. A. multiflorum and A. parviflorum are given a species status. In Turkey, the presence of A. latifolium , A. araxanum , A. lepturoides , A. hohenackerii , A. senganense , A. fominii , A. petraeum , and A. tragacanthinum is also confirmed.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 397–419.