A diploid drop in the tetraploid ocean: hybridization and long-term survival of a singular population of Centaurea weldeniana Rchb. (Asteraceae), a taxon new to Austria

A population of Centaurea weldeniana was discovered recently near Wien (Vienna). The species is reported from Austria for the first time. Determination is confirmed by morphometric analysis of Central European and additional Balkan material of the C. jacea agg. and karyological data. All sampled populations of C. weldeniana are diploid, while only tetraploids have been reported from Central Europe within the C. jacea agg. so far. Detailed morphometric analysis of the newly discovered C. weldeniana population revealed several hybrids with tetraploid C. jacea. The hybrids are tetraploid and originate from unreduced gametes of diploids. No triploids were found. Indeed, the incidence of hybridization is surprisingly low. The population was probably discovered already in 1886 (determined as C. bracteata) and was described as a new species C. argyrolepis in 1901, but these data were lost. However, this isolated diploid population has survived for at least 125?years, which provides a good example of the strength of inter-cytotype reproductive barriers in Centaurea sect. Jacea.     

Distribution ofUrtica dioica L. andRubus fruticosus L. (+agg.) in relation to edaphic factors in cultivated poplar woods

Urtica dioica L. andRubus fruticosus L. (+agg.) are widely distributed in Belgian cultivated poplar woods. Using adjusted standardized residuals as an objective measure of preference of a species for ecoparameters, it seems possible to describe the distribution betweenUrtica andRubus as coincidental as regards ecological considerations. Both plant species behave obviously as antagonists.     

Selected and annotated chromosome counts inTaraxacum sect.Palustria Dahlst. (Studies inTaraxacum 3.)

Chromosome counts of 9 species ofTaraxacum sect.Palustria are given. Five species were karyologically examined for the first time, viz.T. trilobifolium Hudziok,T. lissocarpum agg.,T. paucilobum Hudziok,T. subalpinum Hudziok andT. mendax, a new pentaploid species. Taxonomic and nomenclatural notes are added.     

Origins and distribution of invasive Rubus fruticosus L. agg. (Rosaceae) clones in the Western United States

In facultatively asexual invasive species, an understanding of the origins and diversity of clones can reveal introduction and invasion pathways and inform management efforts. In this study, we use microsatellite and chloroplast DNA markers to infer clonal diversity of the Rubus fruticosus agg. invasion in the Western United States, determine the relationship of these clones to clones found in other exotic ranges, and determine the geographic and genetic origins of the invasive clones. We found two invasive clones in the Western United States, where the invasion had previously been thought to consist of a single asexual lineage. The most common clone was genetically identical to the microspecies R. armeniacus from the native range of Germany, while the second clone was identical to the microspecies R. anglocandicans in the invaded range of Australia and closely related to samples from the native ranges of England and Serbia. A third distinct clone was identified in a collection from the exotic range of Chile. Our results demonstrate that cryptic genetic diversity may be present in asexual invasions that are thought to be homogeneous. However, the asexual relationships between R. fruticosus agg. clones in the native and multiple exotic ranges indicate that preadaptation has played an important role in invasion success in this species aggregate.     

Genetic structure of three Sorbus latifolia (Lam.) Pers. taxa endemic to northern Bavaria

The Franconian Alb (Bavaria, Germany) is rich in endemic Sorbus taxa, considered as apomictic microspecies and derived by hybridization between Sorbus aria aggregate and Sorbus torminalis (Sorbus latifolia aggregate). Molecular studies using the AFLP technique, neighbour joining, Bayesian clustering, principal coordinate analysis (PCo) and voucher studies were used to investigate genetic structure and origin of adult plants and cultivated offspring of three endemic S. latifolia taxa, namely Sorbus cordigastensis, Sorbus franconica and Sorbus adeana and probable parental species from the S. aria agg. and S. torminalis. The S. latifolia taxa, adults and progeny, showed low genetic variability and a more or less clonal structure, confirming an apomictic mode of reproduction. The investigated S. latifolia taxa were remarkably different among each other, confirming their status as microspecies. The AFLP data confirmed the hybrid origin of the S. latifolia taxa, they were 1.3–1.5 times more closely related to S. aria agg. than to S. torminalis. The S. aria agg. showed a complicate genetic structure and fell into four main groups, two intermediate groups besides Sorbus pannonica and Sorbus aria sensu stricto (S. aria s.str.). Some progeny of S. pannonica was more variable than expected and clustered partly with other groups indicating gene flow within S. aria agg. Different subgroups of the S. aria aggregate may be parental for the S. latifolia taxa, contributing to the remarkable genetic distances between them.     

New names inPhanerogamae 5

103 new nomenclatural combinations are proposed for taxa of European flora, 95 for species, 8 for subspecies. Most of them have resulted from a more natural delimitation of genera. The majority of new combinations are made in the segregants ofCentourea (Acosta—16,Colymbada— 8),Hieracium (Pilosella—16),Scabiosa (Trochocephalus—14),Serratula (Klasea—13) andLeontodon (Scorzoneroides—8). Taxonomie or nomenclatural problems of some taxa (Arum orientale, Colobium taraxacoides, Klasea, Leucanthemopsis alpina agg.,Psyllium, Scorzoneroides, Tephroseris cantabrica) are discussed. Names omitted from the Index Kewensis and corrections of publication places are added (17 names).     

Population dynamics of annual Salicornia species in the tidal salt marshes of the Oosterschelde,The Netherlands

1. The population dynamics of two Salicornia species from the Bergen op Zoom salt marsh (south-west Netherlands) was examined. Based on the results of several field studies three preliminary life tables were constructed, two for S. procumbens agg. populations growing respectively on the mud flats and in the salt marsh, and one for S. europaea agg. living in the upper marsh.
2. The life cycles are described and quantified in terms of eight phases and the transition probabilities between them, starting from a notional individual representative of each population.
3. The models depicting the life cycle of S. procumbens show a mean offspring number of 4.26 individuals per parent for the mud-flat population and 0.18 for the salt-marsh population. The S. europaea model gives an output of 0.44 individuals per parent. These results reflect the fluctuations in population size observed in sample plots over the years 1976–78.
4. Comparison of the transition probabilities reveals that on the mud flats most S. procumbens individuals die during pollination and seed germination, while the population in the salt marsh proper is thinned especially during the seed phase in winter time and during the growth from established seedlings to maturation. S. europaea behaves in a similar but less pronounced way to S. procumbens in the salt marsh.
5. Probabilities for one flower or one seed to produce a mature flowering plant were calculated, and were compared with those found in the literature. They are roughly of the same order of magnitude as the probabilities for other annual species, but much higher than those reported for biennial species.

     

Taxonomic individuality of Leonurus cardiaca and Leonurus quinquelobatus in view of morphological and molecular studies

The main goal of this study was to determine the number and taxonomic rank of taxa belonging to the complex Leonurus cardiaca agg. in Poland. Based on statistical analysis of selected features, two morphological forms of this plant were distinguished. In order to determine their genetic polymorphism and the relationships between them, the nuclear, mitochondrial and chloroplast genomes were analysed with the use of RAPD and PCR–RFLP markers. 39 RAPD primers produced a total of 234 nuclear DNA fragments, of which 128 were polymorphic and distributed almost equally between two forms. It was found that 87 % of the compared pairs of RAPD profiles differ from each other. Five chloroplast and two mitochondrial primer pairs were used to amplify non-coding regions of organelle genomes. Restriction analysis revealed uniformity of mtDNA and occurrence of two cpDNA haplotypes, corresponding to naked and hairy forms of L. cardiaca agg. The obtained results justifies the recognition of these forms as separate species L. cardiaca s. s. L. and L. quinquelobatus Gilib., respectively. The distribution of both species in Poland is given in the paper.     

Rubus canduliger, a new regional species from the Netherlands, with notes on the range structure and dynamics of brambles (Rubus, Rosaceae)

Rubus canduliger is described as a new regional bramble species belonging to the seriesDiscolores. It has been recognized in the Netherlands as a distinct species for more than 70 years but identified as the central EuropeanR. grabowskii. The latter, however, differs by its tapering prickles, regular leaves with longer petiolule of the terminal leaflet, broader, often pyramidal inflorescence, and white or pink flowers with carpels hairy at the top.R. canduliger has a well-defined range north of the Rhine river around Arnhem where it occurs in urban and other disturbed habitats. Historical events, e.g. transport of young trees in the 16th century may explain most records south of the Rhine river, 25–50 km outside the main range. The ecology and phytosociology of the new bramble are discussed. Its distribution is compared toR. trichanthus, another regional species with a similar range asR. canduliger, but occurring in more natural habitats. In pseudogamous European blackberries (Rubus sectionRubus =R. fruticosus agg.), range size is considered to be related to the age of the species. Several widespread species have disjunct distribution ranges that are suggested to be the result of long-distance dispersal by migratory birds but unintentional anthropogenic transport of seeds may be involved as well. We hypothesize that in northwestern Europe fragmentation of continuous ranges of widespread species was caused by ongoing deforestation and subsequent degradation (acidification) of originally base-rich sandy and loamy soils from the Iron Age onward. Many regional species, however, have originated and spread in the resulting man-made landscapes. A classification of range types of brambles differing in size and internal structure is presented.     

Chromosome numbers and DNA ploidy levels of selected species ofHieracium s.str. (Asteraceae)

Chromosome numbers and /or ploidy levels are reported for 44 species and subspecies ofHieracium s.str. from the following European countries: Andorra, Austria, Bulgaria, Czech Republic, France, Italy, Montenegro, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland and Ukraine. The chromosome numbers/DNA ploidy levels ofH. bocconei (2n ～ 4x),H. bupleuroides subsp.leviceps (2n = 27),H. caesioides subsp.caesioides (2n = 27),H. basifolium (H. caesium agg., 2n = 36),H. plumbeum (H. caesium agg., 2n = 36),H. glaucum subsp.nipholepium (2n= 27),H.gouanii (2n = 18),H. gymnocerinthe (2n = 27),H. ramondii (2n = 27),H. recoderi (2n = 18),H. stelligerum (2n = 18), andH. tomentosum (2n = 18, 2n ～ 2x, 2n ～ 3x) were determined for the first time. New ploidy levels are reported forH. cerinthoides s.str. (2n = 27),H. humile (2n = 36), andH. tommasinianum (2n = 27).     

Morphological and ploidy level variation ofCentaurea phrygia agg. (Asteraceae) in the Czech Republic, Slovakia and Ukraine

Centaurea phrygia agg. was studied in part of Central Europe (Czech Republic, Slovakia and Ukraine) to evaluate its karyological and morphological variation. Three ploidy levels were found: diploid (C. pseudophrygia, C. stenolepis and several populations ofC. phrygia s.str.), tetraploid (C. oxylepis and several populations ofC. phrygia s.str.), and triploid (very rare, only single individuals). Only one taxon,C. phrygia s.str., comprises populations of two ploidy levels, but they seem to be geographically separated. Multivariate morphometric analysis confirmed the separation of the four taxa; the length and the width of appendages of middle involucral bracts, visibility of appendages of inner involucral bracts, the length and the width of an involucre, and the length/width ratio of middle cauline leaves are the most important discriminant characters. Several populations intermediate between two species (eitherC. pseudophrygia andC. stenolepis orC. oxylepis andC. phrygia s.str.) were found. Their possible origin from introgressive hybridization is discussed considering their morphology (intermediate between putative parental species), chromosome numbers (always the same as in putative parents), and distribution (both spatially and ecologically more or less intermediate between distribution ranges of putative parents). A determination key for the four studied species, including intermediates, is also presented.     

Serpentine and polyploid differentiation withinGalium pumilum agg. (Rubiaceae) in Eastern C. Europe

The local differentiation of a model group,Galium pumilum agg., was studied in serpentine versus non-serpentine habitats in the Czech Republic and adjacent regions in Austria, Germany and Poland. Serpentine habitats form ecologically distinct islands within the landscape; local micro-evolutionary processes may be important there, both because of small population sizes and the strong selection effects of the metal-rich soils. TheGalium pumilum agg. is a group of taxa showing typical karyological and ecological differentiation; the following species and cytotypes were studied:Galium pumilum Murray (2n=8x=88),G. valdepilosum Heinr. Braun (2n=2x=22, 2n=4x=44),G. sudeticum Tausch (2n=4x=44) andG. austriacum Jacq. (2n=4x=44). The karyological and taxonomic investigation showed that serpentine versus non-serpentine habitats do not play an important role in the differentiation of the species studied. (1) The examined populations ofG. pumilum Murray were homogeneous in chromosome numbers, belonging to the octoploid (2n=88) cytotype only. (2)G. valdepilosum consists of two cytotypes. Both of them occur on and off serpentine sites, but they show pronounced East-West distributional difference. (3)G. sudeticum occupies special localities on serpentine in western Bohemia, which are geographically isolated from its nearest localities in the Krkono?e Mts. The plants coming from both localities belong to the tetraploid cytotype. (4) A highly polyploid plant (2n=12x=ca. 132) from theG. pumilum group was found on serpentine. From the methodological point of view, pollen size turned out to be reliable only for distinguishing octoploids from diploids and tetraploids, but it is less suitable for separating the latter two ploidy levels. The occurrence ofG. austriacum is reported for the Czech Republic for the first time with certainty. This record extends the known distribution area ofG. austriacum in the NW direction.     

Chromosome numbers and evolution inOphrys (Orchidaceae)

The diploid basic chromosome number forOphrys is 2n = 36. Tetraploidy has been found inO. fusca agg. only. Aneuploidy and aneusomaty are wide-spread; supernumerary chromosomes behave B-like, but do not differ by shape or structure from A-chromosomes. A new staining method reveals clearly differentiated bipartite meiotic kinetochores. Evolutionary aspects of homogamic hybridization, chromosome instability, and of the basic chromosome number x = 18 are discussed.     

Identification of the agg1 mutation responsible for negative phototaxis in a “wild-type” strain of Chlamydomonas reinhardtii

The unicellular green alga Chlamydomonas reinhardtii is a model organism for various studies in biology. CC-124 is a laboratory strain widely used as a wild type. However, this strain is known to carry agg1 mutation, which causes cells to swim away from the light source (negative phototaxis), in contrast to the cells of other wild-type strains, which swim toward the light source (positive phototaxis). Here we identified the causative gene of agg1 (AGG1) using AFLP-based gene mapping and whole genome next-generation sequencing. This gene encodes a 36-kDa protein containing a Fibronectin type III domain and a CHORD-Sgt1 (CS) domain. The gene product is localized to the cell body and not to flagella or basal body.     

Taxonomic update of Clitocybula sensu lato with a new generic classification

The taxonomy and phylogeny of the hydropoid clade (genera Clitocybula s.l., Megacollybia, Leucoinocybe gen. nov., Hydropus, Trogia, Gerronema, Porotheleum and Lignomphalia gen. nov.) in Europe is studied using morphological and molecular approaches; the first three genera in detail including all known European species. Only two European species remain in Clitocybula s.str., Clitocybula lacerata and Clitocybula familia. The European C. lacerata is a species complex which should be treated as C. lacerata agg. at the current state of knowledge. A neotype originating from type area was designated to fix the application of the name. The presence of American species Clitocybula abundans in Europe is insufficiently proved. “Clitocybula dryadicola ? belongs to the genus Hydropus, and Clitocybula tilieti has an unclear systematic position. The results showed that Megacollybia and Leucoinocybe represent independent genera separated from Clitocybula. The genus Leucoinocybe is validly published with two European species, Leucoinocybe lenta and Leucoinocybe taniae. “Clitocybula flavoaurantia” proved to be conspecific with the latter species. The genus Lignomphalia is published for “Pseudoomphalina lignicola”, a lignicolous omphalinoid species. The Indian “Clitocybula sulcata” is transferred to Leucoinocybe and “Clitocybula atrialba” to Gerronema. The first European records of Megacollybia marginata are published.     

AFLP analyses demonstrate genetic divergence, hybridization, and multiple polyploidization in the evolution of Achillea (Asteraceae-Anthemideae)

Achillea, a temperate genus of herbaceous allogamous perennials, is a model for evolutionary radiation through hybridization and polyploidization. AFLP analyses were performed on 300 individuals of 66 populations and 27 taxa/cytotypes, mainly from the polyploid A. millefolium aggregate and its suspected hybrid links with other clades of the genus. The mosaic genetic structure of hybrids and polyploids is revealed by specific AFLP bands shared with their assumed parents. In E Asia, A. alpina-4x and A. wilsoniana-4x are allotetraploids between A. acuminata-2x (sect. Ptarmica) and A. asiatica-2x (sect. Achillea-A. millefolium agg.). A. virescens-4x is a hybrid species linking A. nobilis agg. and A. millefolium agg. in S Europe. The hybrid swarm A. clypeolata-2x yen A. collina-4x recently formed in Bulgaria shows no AFLP bands additive to its parents; by contrast, other more ancient allopolyploids exhibit genetic innovations. Relationships within A. millefolium agg. are complex. Five 2x-taxa, mostly well separated and regressive, are limited to Eurasia; seven 4x- and 6x-taxa are intimately linked by hybridization, are expansive, and through A. asiatica-2x/4x have formed the N American polyploids. All these results from AFLPs correspond well to other evidence, and indicate a long history of reticulate evolution in Achillea.     

Festuca austrodolomitica,a new species of theF. halleri group (Poaceae) from the SE Alps

The new species belongs toFestuca halleri agg. (F. ovina s. latiss.) and is diploid. Its habitat are alpine snow bed communities (Arabidetalia caeruleae) of the southern Alps, between the Brenta and Mt Cadria in the W and Mt Cavallo near Belluno in the E. Morphological, anatomical, epidermal, and ecological comparisons with related species suggest that it is closest to the vicariousF. rupicaprina in the N Alps.     

Contribution to the biology of germination of four species ofChenopodium album agg. under different condition

The study investigated various factors stimulating germination of seeds of four species ofChenopodium album agg.—C. album L. s. str.,C. ficifolium Sm.,C. strictum Roth,C. suecicum J. Murr. These factors were: storage in wet and cold, alternating temperature, nitrates mainly in combination with light and, to a lesser degree, the duration of chilling. Germination was inhibited by darkness, which was most marked with seeds ofC. strictum. The effect of alternating temperature was better on the germination ofC. ficifolium andC. suecicum than on that ofC. album andC. strictum. We also discussed several differences in specific reactions of the individual species to the factors under consideration.     

Challenges in the delimitation of morphologically similar species: a case study of <Emphasis Type="Italic">Tuber brumale</Emphasis> agg. (Ascomycota,Pezizales)

Tuber brumale (winter truffle) is one of the most controversial true truffles, not only in regard to its ecological and economical role but also its taxonomy. Multilocus phylogenetic analyses have revealed that specimens identified earlier as T. brumale belong to two species. These species were deemed cryptic right away, because preliminary morphological measurements did not show any phenotypical differences. In this study, we measured the morphology of 119 T. brumale agg. specimens, identified by DNA-based phylogenetic tools. We found several continuous morphological characters which show strong statistical differences between the two species, albeit not without overlap. Using a combination of these characters, we show that efficient separation of the two species is possible. We describe T. cryptobrumale sp. nov. and present the environmental demands and the potential area reconstruction of both species. We argue that non-representative sampling is a major culprit in most failures to detect both the existence of morphologically similar species and their morphological differences. Our findings illustrate the benefits of integrative taxonomy: the use of a combination of molecular, morphological and ecological tools.     

Die Chromosomenzahlen von tschechoslowakischen Arten der GattungKnautia L. (Dipsacaceae)

Die Chromosomenzahlen von den tschechoslowakischen Sippen der GattungKnautia L. sind angegeben:K. drymeia Heuffel subsp.drymeia—2n = 40 (41,42),K. slovaca ?těpánek— 2n = 20,K. arvensis (L.)Coulter subsp.arvensis—2n = 20, 40 (39, 41),K. arvensis (L.)Coulter s. 1.—2n = 20,K. kitaibelii (Schult.) Borbás—2n = 20 (39, 41),K. dipsacifolia Kreutzer subsp.dipsacifolia—2n = 60,K. dipsacifolia agg.—2n = 40, 50, 60. Keine taxonomisch wertvolle morphologische Unterschiede zwischen diploiden und tetraploiden Zytotypen vonK. arvensis subsp.arvensis wurden gefunden. Für die Populationen von Pflanzen, deren Morphologierauf Kreuzung oder Introgression zwischen tetraploiden Sippen (K. arvensis subsp.arvensis—4x,K. drymeia subsp.drymeia, K. kitaibelii undK. dipsacifolia agg.) zeigt, wurden ím Einklang mit dieser Voraussetzung die tetraploiden Chromosomenzahlen—2n = 40 (39, 41)— festgestellt. Einige ungelöste taxonomische Fragen hauptsächlich über die diploiden Populationen vonK. arvensis s. l. in Böhmen und überK. dipsacifolia agg. in den Westkarpaten werden diskutiert.