A revision ofTaraxacum sect.Piesis (Compositae)

On the basis of allozyme and cultivation data, and of additional herbarium material, a taxonomic and nomenclatural revision ofTaraxacum sect.Piesis A.J. Richards exKirschner et?těpánek is provided. The section is made up of halophilous, sexually reproducing taxa. InT. stenocephalum Boiss. etKotschy,T. pindicum Kirschner et?těpánek, sp. nov., andT. perenne Kirschner et?těpánek, sp. nov., a tetraploid chromosome number has been recorded, representing the only known case of sexuality at the tetraploid level in the genus. The complex ofT. stenocephalum, includes some geographically and morphologically extreme populations treated as subspecies: subsp.gumusanicum (Soest)Kirschner et?těpánek, comb. nov., subsp.magnum Kirschner et?těpánek, subsp. nov., and subsp.daralagesicum (Schischk.)Kirschner et?těpánek, comb. nov. In addition toT. bessarabicum (Hornem.)Hand.-Mazz., a widely distributed Eurasian species,T. stenocephalum, a complex centred in Transcaucasia and Anatolia, andT. pachypodum H. Lindb., a North African endemic, four new species are described:T. salsum Kirschner et?těpánek, sp. nov., a diploid endemic confined to E Crimea,T. perenne Kirschner et?těpánek, sp. nov., a tetraploid sexual species known only from SW Crimea,T. pindicum Kirschner et?těpánek, sp. nov., a remarkable tetraploid endemic to the Pindos Mts., Greece, andT. salsitatis Kirschner, ?těpänek etYirdirimli, sp. nov., an Anatolian diploid species. Furthermore, a hybrid betweenT. salsum andT. bessarabicum from Crimea (documented on the basis of allozyme data elsewhere) is given a binomial,T. xmesohalobium Kirschner et?těpánek, nothosp. nov.     

Allozyme divergence among species ofWolffia (Lemnaceae)

The genusWolffia was surveyed electrophoretically at 14 allozyme loci. A total of 133 clones representing 10 of the 11 recognized species was examined. Genetic identities among most pairs of species are zero, with non-zero values ranging from 0.14 to 0.40.Wolffia angusta and the newly describedW. neglecta show the highest similarity, and the former species has an identity of 0.14 withW. australiana. The next highest similarity (0.34) occurs betweenW. globosa of Southeast Asia andW. cylindracea of southern Africa, which until recently, had generally been viewed as members of the same species. Other species showing some common alleles are members of a complex involvingW. arrhiza, W. columbiana, W. cylindracea, andW. globosa. WithinW. arrhiza, plants from South Africa and Europe are easily distinguished electrophoretically because each contains unique alleles at two loci. Strains from other parts of Africa vary at these loci and are not totally distinct from either the plants from South Africa or from Europe. Species ofWolffia are much more divergent at allozyme loci than the majority of congeners of flowering plants. This suggests that the species are quite old and that the difficulties in distinguishing taxa morphologically are the result of reduction rather than lack of divergence due to recent speciation. Because of the lack of shared alleles between the majority of species pairs inWolffia, enzyme electrophoresis provides limited resolution of species relationships in the genus.     

Allozyme analysis ofLuzula sect.Luzula (Juncaceae) in Ireland: Evidence of the origin of tetraploids

Luzula campestris, L. pallidula and tetraploi plants of this section from Ireland were studied electrophoretically. A hypothesis of alloploid origin for the Irish tetraploids from a hybrid betweenL. campestris andL. pallidula, based on the morphology, and karyology of the relevant taxa, has been tested. The presumed parental taxa are divergent in several loci,L. pallidula being highly homozygous andL. campestris moderately heterozygous. The Irish tetraploid combines the alleles of the putative parents, has no unique alleles, and exhibits a high frequency of heterozygotes, inAdh-2 andGpi-1, with invariably additive patterns. The Irish tetraploid has a low genotype diversity. The allozyme data support the hypothesis of alloploid origin for the Irish tetraploid withL. campestris andL. pallidula as possible parents.     

Relationships between species ofLeymus,Psathyrostachys, andHordeum (Poaceae,Triticeae) inferred from Southern hybridization of genomic and cloned DNA probes

We have used total genomic DNA as a probe to size-fractionated restriction enzyme digests of genomic DNA from a range ofTriticeae species from the generaLeymus Hochst.,Psathyrostachys Nevski, andHordeum L., and hybrids betweenHordeum andLeymus to investigate their taxonomic relationships. Genomic Southern hybridization was found to be an effective and simple way to assess the distribution and diversity of essentially species-specific and common, repetitive DNA sequences, and is hence especially useful in evolutionary studies. The DNA sequences ofH. vulgare seem to diverge substantially from those ofH. brachyantherum, H. lechleri, H. procerum, andH. depressum. The genome ofThinopyron bessarabicum shows little homology to those of theLeymus species investigated, confirming thatT. bessarabicum is not an ancestral genome inLeymus. Although the genomes ofLeymus andPsathyrostachys share substantial proportions of DNA sequences, they include divergent repeated sequences as well. Hybridization with a ribosomal DNA probe (pTa 71) showed that the coding regions containing structural genes encoding the 18 S, 5.8 S, and 26 S ribosomal RNA were conserved among the species investigated, whereas the intergenic spacer region was more variable, presenting different sizes of restriction fragments and enabling a classification of the species. The rye heterochromatin probe pSc 119.2 hybridized to DNA fromH. lechleri andT. bessarabicum, but not to DNA from the other species investigated.     

Ontogenetic variations of the Early Cretaceous non-marine bivalveTrigonioides (T.) heilongjiangensis and the concept of fossil species

With growth, the angle of V-ribs on the median flank ofTrigonioides (T.) heilongjiangensis enlarges; the subradial ribs on the anterior half vary from simple, but prominent, through feeble or vanished to bifurcate or multiple and clear; the number of submedian teeth increases and the variation in outline of the species is quite continuous. All individuals or populations which vary continuously in characters at one time horizon and/or in a certain time space within the same biogeographic region should be regarded as conspecific. Accordingly, the individuals of five taxa ofTrigonioides hitherto recognized in the eastern Heilongjiang area, namelyT. (T.) kodairai Kobayashi & Suzuki,T. (Wakinoa) wakinoensis intermedia Hase,T. (W.) subtrigonus Sha,T. (W.) cf.tamuraia Yang andTrigonioides sp., are regarded as representing various ontogenetic stages ofTrigonioides (T.) heilongjiangensis Sha & Gu. 263     

Isozymes, and the status of Taraxacum (Asteraceae) agamospecies

HUGHES, L. & RICHARDS, A. J., 1989. Isozymes and the status of Taraxacum (Asteraceae) agamospecies. Genetic identities I and Similarity Indices SI are calculated between 12 samples of Taraxacum, on the basis of 40 isozymes at 15 loci for 10 enzyme systems. Samples included three polyploid agamospermous populations from northern England (group 1), three sexual diploid populations from south-central France (group 2), and six accessions of ‘primitive’ diploid self-compatible sexual taxa from southern Europe and Australia. Samples could be assigned to eight species, classified in seven sections of the genus. Two clusters of high relationship were evident. All the group 1 and group 2 species were very closely related, with pairwise comparisons for I in excess of 0.93. The three group 3 accessions identified as T. bessarabicum showed pairwise comparisons for SI in excess of 0.71. Comparisons for SI between the other group 3 species, and between all the group 3 species and the group 1 and 2 species were all very low, not exceeding 0.45. It is concluded that dissimilarity between samples as assessed by isozymes is probably related to the time of evolutionary divergence of those samples. Although allopolyploid, and morphologically very diverse, the group 1 agamospecies may have very recently diverged asexually from a common stock. The group 2 diploids may have resulted from rediploidization and regained sexuality from the same originally agamospermous stock. In areas of Europe in which such ‘modern’ sexuality is common, it is probable that all ‘modern’ Taraxaca, including at least five sections of the genus, should be included within a single taxon. In contrast, ‘primitive’ self-compatible sexual species in group 3 appear to have diverged from each other several million years ago, and with the exception of the disjunct accessions of T. bessarabicum, are genetically highly distinct. Such species should be maintained in the taxonomies of all areas. It is probable that an agamospecies classification of ‘modern’Taraxacum will continue to convey much useful information in areas, such as northern Europe, in which sexuality is absent.     

Tetraploids inLuzula multiflora (Juncaceae) in Ireland: Karyology and meiotic behaviour

Populations ofLuzula multiflora s.l. in Ireland were examined karyologically. Plants from 14 populations were invariably tetraploid with 2n=24. Chromosomes of the tetraploid are of AL type (true tetraploidy). Meiosis of the tetraploids is of the same type as described for otherLuzula taxa in the literature. In meiosis, 12 bivalents are regularly formed. A hypothesis based on the morphological and allozyme data, that the tetraploids are of alloploid origin, is supported by the present results. Meiosis in an artificial hybrid between the presumed parental taxa,L. campestris andL. pallidula, was studied; a certain tendency towards chromosome doubling was observed. The geographical distribution of theL. multiflora cytotypes is also discussed.     

Genetic diversities of four little-known species of Malesherbia (Malesherbiaceae) endemic to the arid inter-Andean valleys of Peru

Few studies of genetic variation have been conducted on plants of the Pacific coastal desert and neighboring Andes of Peru, although the region has many endemic taxa. Enzyme electrophoresis was employed to examine allozyme diversities of four species of the family Malesher-biaceae, an endemic to the arid Andes and coastal desert. One population ofMalesherbia splendens and two ofM. tubulosa, both endemics of the department of Lima, one population ofM. weberbaueri var.weberbaueri, an endemic of the Andean department Huancavélica, and the two Lima populations ofM. scarlatiflora were studied. Fifteen loci were examined for all populations and an additional seven loci were resolved forM. weberbaueri andM. splendens. Malesherbia splendens, which is known from three populations, has a low mean number of alleles per locus (A), proportion of polymorphic loci (P), and expected heterozygosity (H_s) (A=1.214, P=0.214, H_s=0.057).Malesherbia weberbaueri (A=1.231, P=0.154, H_s=0.079) andM. scarlatiflora (A=1.364, P=0.273, H_t=0.083) both have average expected heterozygosities and relatively low mean numbers of alleles per locus and proportions of polymorphic loci. InM. tubulosa, all measures of genetic diversity are high in comparison with other endemics (A=1.818, P=0.364, H_t=0.206).Malesherbia tubulosa has high interpopulation differentiation, whereasM. scarlatiflora has low among-population diversity. The relatively low allozyme diversities, restricted habitats, narrow ranges ofM. splendens andM. weberbaueri, and the morphological similarities of all four species suggest that they evolved recently by founder events. Greater allozyme diversities inM. tubulosa could be attributable to its maintenance of larger populations in a greater variety of habitats.     

Taxonomic studies of theMuscari botryoides complex in Hungary

The ploidy level and karyotype ofMuscari botryoides (s.l.) samples from Hungary (25 localities) and Romania (1 locality:locus classicus ofM. transsilvanicum) were determined. The Romanian sample proved to be diploid (2n=18), while in Hungary both diploid and tetraploid (2n=36) populations occurred. The karyotypes of all diploid populations were similar: 2 pairs of long acrocentric (one of them usually with satellites) + 3 pairs of medium-sized submetacentric-metacentric + 4 pairs of short ± metacentric chromosomes. All diploid populations in Hungary can be identified asM. transsilvanicum. There is no reason to support the taxonM. botryoides subsp.hungaricum because it does not differ from the sample collected at thelocus classicus ofM. transsilvanicum (Romania, Sibiu-Gu?teri?a) in any of the characteristics mentioned in its protologue. Its karyotype also corresponds to that ofM. transsilvanicum. Contrary to the former assumptions, the tetraploidM. botryoides is also native to Hungary. The tetraploid karyotype seems to be somewhat of a duplication of the diploid one. Morphological characters used in the identification keys are not suitable for unambiguous separation of the taxa mentioned above, though morphometric analyses revealed some quantitative differences between diploids and tetraploids. Their separation on species level can only be supported by the supposed reproductive barriers caused by different ploidy level and chorology. In HungaryM. transsilvanicum is restricted mostly to the Eupannonian Region, the Mecsek and Villány Mts.M. botryoides does not occur in the Eupannonicum, instead it inhabits the subatlantic hilly W and SW part of Hungary and the Northern Mountain Range. The latter territory (including also the Slovak localities) seems to be the easternmost extension of the area ofM. botryoides.     

Speciation inVirgilia (Fabaceae): Allopatric divergence followed by introgression?

Levels of variation revealed by starch gel-electrophoresis were compared with morphological and chemical variation within and among the species and subspecies ofVirgilia:V. oroboides subsp.oroboides, V. oroboides subsp.ferruginea andV. divaricata. The data sets exhibited concordance in that all point to a very close relationship between the taxa. Differences are mostly quantitative and an overlapping east-west gradient of character variation is indicated. Analysis of morphological and chemical characters showed thatV. oroboides subsp.ferruginea andV. divaricata are relatively distinct, whereas allozyme analysis indicated a high degree of genetic similarity among populations of these two taxa. The observed pattern of variation suggests relatively recent speciation with subsequent introgressive hybridisation resulting in a geographical and ecological gradient.     

Taxonomy of theHieracium alpinum group in the sudeten Mts., the West and the Ukrainian East Carpathians

A taxonomic revision of theHieracium alpinum group (sensu Flora Europaea) in the Sudeten Mts., the West and the Ukrainian East Carpathians is provided. Six species are distinguished in the area studied, viz.Hieracium alpinum, H. halleri, H. augusti-bayeri, H. melanocephalum, H. tubulosum, andH. schustleri. H. alpinum occurs throughout the mountain ranges studied, three other taxa are endemic to the Sudeten Mts. (H. melanocephalum, H. tubulosum, H. schustleri), one is confined to the West Carpathians (H. halleri, in addition to the Alps) and one is endemic to the Ukrainian East Carpathians (H. augusti-bayeri). A diploid cytotype (2n=18) was ascertained inH. alpinum from the Ukrainian East Carpathians and the same chromosome number was found forH. augusti-bayeri. An agamospermous mode of reproduction was confirmed for the triploid cytotype ofH. alpinum, the triploidH. melanocephalum andH. halleri and the tetraploidsH. tubulosum andH. schustleri; the diploid taxa were found to be sexual. Pollen production in diploid taxa is high and pollen grains are homogeneous in size; triploid species, as well as tetraploidH. tubulosum, do not produce polllen; tetraploidH. schustleri has pollen grains of variable size. Data on the ecology and distribution of the species are also given.     

Three new species and a new combination in the genusTorulomyces from soil

Seven isolates ofTorulomyces from Asian and Australian soil samples were studied in comparison with known taxa of the genus and withMonocillium indicum, the type species ofMonocillium. Three new species,Torulomyces parviverrucosus, T. laevis, andT. ovatus, are described, andT. brunneus is described as a new combination. Conidial characteristics, especially their shape and surface structure, are useful taxonomic criteria for distinguishing species ofTorulomyces. Monocillium is considered to be a distinct genus.     

Reclassifications and new names for some European Phanerogams

On the basis of taxonomic and nomenclatural revisions of various plant groups 37 new nomenclatural combinations are proposed mostly for European phanerogamous taxa (21 for species, 14 for subspecies, 2 for hybrids.)Medicula Medicus,Pseudofumaria Medicus andThlasidia Rafin. are accepted as taxonomically justified genera and correct generic names.Dactylorhiza longebracteata (F. W. Schmidt)Holub is proposed instead ofD. fuchsii (Druce) Soó, andHylotelephium triphyllum (Haworth) Holub isstead ofH. purpureum (L.)Holub.     

Allozyme diversity and systematics inAnnonaceae—a pilot project

Five species ofAnnona and one species fromArtabotrys, Cananga, Polyalthia, andRollinia were investigated in regard to 11 different allozyme loci. Preliminary studies on small population samples ofAnnona suggest genetic uniformity in three species and variability within and between populations in two other species. The allotetraploid origin ofA. glabra is clearly shown by its hybrid enzyme bands. The genetic distance between fiveAnnona species partly corresponds with their morphological relationships; onlyA. muricata appears more separated than is suggested by morphology. A comparison of the five genera demonstrates close relationship betweenAnnona andRollinia. Two enzyme loci are identical within all taxa investigated and possibly may serve as a genetic marker for the family.     

Thinopyrum bessarabicum: biochemical and cytological markers for the detection of genetic introgression in its hybrid derivatives with Triticum aestivum L.

Summary Thinopyrum bessarabicum (2n = 2x = 14, JJ) with its unique property of salt tolerance provides a potential means for the transfer of this important and complex trait into cultivated wheat through intergeneric hybridization. To accomplish this, diagnostic markers for detecting the presence of Th. bessarabicum chromosomes in a wheat background have to be established. The C-banded karyotype of Th. bessarabicum distinctly identifies individual Th. bessarabicum chromosomes and separates them from those of Triticum aestivum. Also, seven protein/isozymes, i.e., malate dehydrogenase, high-molecular-weight glutenin, Superoxide dismutase, grain esterase, glutamate oxaloacetate transaminase, -amylase and -amylase, were identified as being positive markers specific to Th. bessarabicum; these were also expressed in the T. aestivum/Th. bessarabicum amphiploid. These diagnostic biochemical markers could be useful in detecting and establishing homoeology of Th. bessarabicum chromosomes in T. aestivum/Th. bessarabicum intergeneric hybrid derivatives.     

Biochemical and molecular diagnostics of Thinopyrum bessarabicum chromosomes in Triticum aestivum germ plasm

Thinopyrum bessarabicum (2n=2x=14, JJ) is a self-fertile salt-tolerant grass species, and its hybridization with Triticum aestivum to achieve the transfer of this attributes has been promoted. For the detection of alien introgression, development of diagnostic markers of Th. bessarabicum chromosomes in the wheat background has emerged as an important aspect in our intergeneric hybridization program. Six proteins/isozymes-high-molecular-weight glutenins, superoxide dismutase, grain esterase, β-amylase, glutamate oxaloacetate transaminase and α-amylase —were identified as positive markers for detecting the presence of Th. bessarabicum chromosomes in the advanced backcross derivatives of T. aestivum/Th. bessarabicum//n^* T. aestivum. Fluorescent in situ hybridization further enabled the detection of complete and translocated arms of Th. bessarabicum chromosomes in the T. aestivum background. These diagnostic markers served for tentatively characterizing a distinct set of Th. bessarabicum disomic additions to wheat (2n=44) and have facilitated establishing the homoeology of these added chromosomes.     

Silurian (Ludlow) acastid trilobites from Gotland and Scania,Sweden

Acastid trilobites from Ludlow strata on Gotland include three stratigraphically non-overlapping species ofAcastella Reed:Acastella madidipes n. sp. (Hemse Marl and Eke Beds),A. breviceps (Angelin 1851) (Burgsvik Beds), andA. amatrix n. sp. (Hamra Beds, and probably Burgsvik Beds). Type and other material ofA. breviceps (Angelin) from the Klinta Formation, Scania, is revised. Meraspid and early holaspid material ofAcastella is described. Some characters supportAcaste and allied taxa of mostly Wenlock age as a monophyletic subfamily Acastinae.Acastella as widely recognized is a grade with Devonian species more closely related to Asteropyginae than to the Ludlow type species. Baltic Ludlow-P?00ED;dolí species previously placed inScotiella Delo form a distinct group,Ewacaste n. gen., that includes a species from the uppermost Eke Beds on Gotland.     

Phylogenetic conclusions from Giemsa banding and NOR staining in top onions (Liliaceae)

The chromosomes of different strains of the top or tree onion, ofAllium cepa andA. fistulosum, as well as of cloned progenies from reciprocal crosses between these two taxa have been studied by application of Feulgen- or aceto carmine-, Giemsa- and silver staining. It was possible to differentiate between the satellite chromosomes and 2–4 other chromosome pairs ofA. cepa andA. fistulosum. The phylogenetic origin of the top onions [A. ×proliferum (Moench)Schrad.] from hybridization ofA. cepa andA. fistulosum is substantiated, taking into consideration the variability in size and position of satellites and of active NORs.     

Genetic differentiation ofOryzias minutillus in Thailand

An allozyme survey was conducted in eleven local populations ofOryzias minutillus from Thailand. The region-specific distribution of alleles and the genetic relationship among the eleven populations revealed that they represented major three population groups, within the country as a whole; the Peninsular, Mae Nam Chao Phraya and Mekong subpopulations. Because their distribution boundaries coincide with geographic features, it is supposed that their genetic differentiation is primarily due to geographic isolation. Karyotype polymorphism has been reported only from the Mae Nam Chao Phraya subpopulation, suggesting that the variant karyotype evolved after allopatric isolation of the three subpopulations.     

Karyological studies in IberianSonchus (Asteraceae: Lactuceae): S. oleraceus, S. microcephalus andS. asper and a general discussion

Chromosome numbers and karyotypes ofSonchus oleraceus (n=16, 2n=32),S. asper (n=9, 2n=18 for subsp.asper and subsp.glaucescens, andS. microcephalus (n=15, 2n=30) are studied; for the first time their idiograms are given. Karyotypes mainly comprise small chromosomes with a degree of asymmetry 2B inS. oleraceus andS. microcephalus and 2A or 2A-2B inS. asper. No karyological differences can be pointed out betweenS. asper subspecies. Data presented support the amphiploid character ofS. oleraceus, and the origin ofS. microcephalus through a dysploid process involving the former taxon. In Iberian representatives of the genus, diagrams of karyotype asymmetry indices show a cluster grouping for species, with the detached exceptions ofS. maritimus andS. crassifolius, which spontaneously hybridize in central Spain. A review of available karyological data shows that in the evolution of the genusSonchus s.l. and relatives, the basic chromosome number x=9 has generally been maintained. Dysploidy is restricted to the seriesS. bourgeaui (n=8) —S. tenerrimus (n=7) andS. oleraceus (n=16) —S. microcephalus (n=15), evolutionarily related and included in the present subgenusSonchus. Polyploidy has been detected in a total of nine taxa ofSonchus and in the generaEmbergeria, Kirkianella, andDendroseris, being more common in peripheral regions of the distribution area of the group. Five diversification centers are proposed for the whole group, of which the Western Mediterranean area, including the Iberian Peninsula, is related to diversification of the present subgenusSonchus.