Chloroplast DNA variation inDesmodium subgenusPodocarpium (Leguminosae): Infrageneric phylogeny and infraspecific variations

Chloroplast DNA (cpDNA) restriction site variation was examined in five species ofDesmodium subgenusPodocarpium (Leguminosae; Papilionoideae; Desmodieae). Twenty four phylogenetically informative cpDNA mutations were scored. The cladistic analysis of characters based on the 24 mutations resulted in the most parsimonious tree which supports the monophyly of the subgenus.Desmodium elegans of subgenusDollinera was the sister group of subgenusPodocarpium in this tree. The groupings obtained from the cpDNA characters were consistent with the present infrageneric classification system for the subgenus except for the infraspecific taxa ofD. podocarpum. Three groups withinD. podocarpum, which were incongruent with the infraspecific classification of the species, were distinguished by a total of four site mutations. The first group consisted of subsp.podocarpum, subsp.fallax, and subsp.oxyphyllum var.oxyphyllum; the second subsp.oxyphyllum var.oxyphyllum; and the last subsp.oxyphyllum var.oxyphyllum and var.mandshuricum.     

Distinction in morphology and esterase isozyme betweenEupatorium glehni (=E. chinense subsp.sachalinense) andE. chinense var.oppositifolium (compositae)

The taxonomic status ofEupatorium chinese var.oppositofolium andE. glehni (=E. chinense subsp.sachalinense) inE. chinense complex semsu Kitamura has long been controversial. In this paper, the degree of divergence between diploids of these two taxa was examined by means of morphological studies including principal component analysis, the electrophoretic analysis of esterase isozyme variation and observations on habitats. The data obtained through the examinations indicate these two taxa are diverged enough to be recognized as distinct biological species. Since the polyploidE. chinense var.oppositifolium is more or less intermediate in morphology between the two diploid taxa, it is considered to have masked the distinction between the two diploid taxa. Also, electrophoretic evidence suggests that polyploidE. chinense var.oppositifolium is not a hybrid or hybrid derivative withE. glehni as a parental species. Possible origin of polyploidE. chinense varoppositifolium is also discussed.     

Chloroplast DNA evidence for introgression and long distance dispersal in the desert annualSenecio flavus (Asteraceae)

Phylogenetic analysis of chloroplast DNA (cpDNA) restriction site variation supports a close genetic relationship between the Southwest AsianSenecio flavus subsp.breviflorus and the North AmericanS. mohavensis. The intercontinental disjunct distribution of these two desert annuals may have originated via long distance dispersal. The chloroplast genomes of the Southern and North AfricanS. flavus subsp.flavus and subsp.breviflorus differ by at least ten restriction sites, while at most two restriction sites differentiate the cpDNA genomes of subsp.breviflorus and the outgroupS. squalidus. This suggests that the cpDNA genome ofS. flavus subsp.breviflorus may have resulted from introgression and chloroplast capture with a Mediterranean species related toS. squalidus. This hypothesized introgression could account for the morphological distinctiveness and duplicated isozyme loci ofS. flavus subsp.breviflorus relative to subsp.flavus.     

Isozyme evidence and the origin ofSenecio vulgaris (Compositae)

An electrophoretic survey of isozyme variation was conducted to test the hypothesis thatSenecio vulgaris L. (2n = 40) is of autotetraploid origin fromS. vernalis Waldst. & Kit. (2n = 20). It was established thatS. vulgaris exhibited fixed heterozygosity at three loci examined, showed disomic inheritance at all polymorphic loci, and contained a gene (Est-1) and an allele (Aat-3b) which were not present in the single population ofS. vernalis surveyed. From this it is concluded thatS. vulgaris is not of autotetraploid origin. Instead, the genetic evidence is in keeping with an allopolyploid origin ofS. vulgaris with the possibility thatS. vernalis acted as one of its two parents.     

Protecting evolutionary significant units for the remnant populations of <Emphasis Type="Italic">Berchemiella wilsonii</Emphasis> var. <Emphasis Type="Italic">pubipetiolata</Emphasis> (Rhamnaceae)

Berchemiella wilsonii var. pubipetiolata (Rhamnaceae) is an endangered tree in eastern China. Habitat destruction has resulted in fragmentation of remnant populations and extinction of local populations. AFLP and cpDNA markers were used to determine the population structure of remnant populations of B. wilsonii var. pubipetiolata. Moderate nuclear genomic diversity was found within each of the four remnant populations (H _S = 0.141–0.172), while the cpDNA haplotype diversity in each population ranged from 0.356 to 0.681. Six haplotypes were identified by a combined cpRFLP and cpSSR analysis in a total of 89 individuals. AMOVA revealed significantly AFLP genetic differentiation within and between regions (Φ_SC = 0.196, Φ_CT = 0.396, respectively), and a high cpDNA haplotype differentiation between regions (Φ_CT = 0.849). The results suggest low gene flow between populations of B. wilsonii var. pubipetiolata. Strong genetic divergence between two regional populations as revealed by both AFLP and cpDNA markers provided convincing evidence that two distinct evolutionary lineages existed, and should be recognized as ‘evolutionary significant units’ (ESUs) for conservation concerns.     

The allopolyploid origin ofSedum rupestre subsp.rupestre (Crassulaceae)

InSedum rupestre L. a polyploid series (x = 16) occurs in which aneuploid chromosome numbers and odd levels of ploidy prevail. The most common and widely distributed cytotype,S. rupestre subsp.rupestre, is 2n = 112. Plants resemblingS. rupestre subsp.rupestre can be obtained by hybridizing the tetraploid cytotypes ofS. forsterianum Sm. (2n = 48) andS. rupestre subsp.erectum 't Hart (2n = 64). Comparison of these artificial hybrids with their parents and a large number of plants ofS. rupestre subsp.rupestre (2n = 112) from nature showed thatS. rupestre subsp.rupestre and the artificial hybrids are morphologically indistinguishable, and intermediate betweenS. forsterianum andS. rupestre subsp.erectum. MorphologicallyS. rupestre subsp.rupestre is closer to subsp.erectum than toS. forsterianum. Chloroplast DNA restriction patterns ofS. rupestre subsp.rupestre, however, resembleS. forsterianum more closely. The combined results of the hybridization experiments, the analysis of the cpDNA restriction patterns, and the morphological variation indicate the allopolyploid origin ofS. rupestre subsp.rupestre. Natural hybrids inSedum (Crassulaceae) 4.     

Isoenzyme evidence on the systematics ofHordeum sectionMarina (Poaceae)

Morphological differentiation of diploid accessions ofHordeum marinum Huds. s.l. into two varieties, var.marinum and var.fouilladei (Rouy)Nevski is associated with isoenzyme differentiation. The tetraploid form ofH. marinum s.l. exhibited fixed heterozygosity of several heterozymes with one homoeozyme shared with var.fouilladei and the second homoeozyme not found in the two diploids. It also differed from both diploids in the mobility of glucose-6-phosphate dehydrogenase. All three taxa differed in morphs of EST-A. It is concluded that the tetraploid is an allopolyploid with one genome closely related to the diploid var.fouilladei and with the second genome divergent from those of both diploids by genes for unique morphs of eight homoeozymes. On the basis of the isoenzyme data, three phylogenetic sibling species—H. marinum Huds. s.str. (2x),H. geniculatum All. s.str. (= var.fouilladei, 2x), andH. caudatum Jaaska, spec. nova (4x), are proposed within theH. marinum s.l. complex and a key is given.     

Quantitative assessment of heteroplasmy levels in <Emphasis Type="Italic">Senecio vulgaris</Emphasis> chloroplast DNA

Heteroplasmy in coding chloroplast DNA was only recently shown to occur and was so far not quantitatively assessed. We present a quantitative analysis of cpDNA heteroplasmy levels at a triazine-resistance determining site within and between individual Senecio vulgaris plants. Detectable levels of heteroplasmic haplotypes were observed in all tested plants. As expected, the levels of heteroplasmy vary greatly between plants. However, even within individual plants, the fraction of one haplotype may cover a range from below 1 to well over 90. Our results suggest that heteroplasmy may be a common phenomenon in S. vulgaris.Possible consequences for molecular diagnostics of chloroplast encoded traits as well as evolutionary consequences of chloroplast heteroplasmy are discussed.     

Restriction fragment length polymorphisms of the phytohemagglutinin genes inPhaseolus andVigna (Leguminosae)

Restriction fragment length polymorphisms at the phytohemagglutinin (PHA) locus were determined among 21 genotypes ofPhaseolus vulgaris, P. coccineus, P. acutifolius, P. lunatus, and threeVigna species, using five restriction enzymes and one double digestion, in order to provide molecular evidence for their genetic relatedness. The dissimilarity between genotypes was estimated from binary RFLP data. The dissimilarity was high among species (from 0.75 to 0.95), and of variable extent among genotypes of the same species (0.33–0.89). InP. vulgaris, two different DNA hybridization patterns were found, giving further evidence for two major gene pools in that species. The restriction patterns ofP. vulgaris var.aborigineus, the putative ancestral form ofP. vulgaris, exhibit clear homology toP. vulgaris genotypes. An undefined landrace from Taiwan could be identified as aP. vulgaris genotype. RFLP-based trees for the phytohemagglutinin genes of the species studied were computed with several distance matrix and parsimony methods.     

Giemsa karyotypes and their evolutionary significance inScilla bifolia,S. drunensis,andS. vindobonensis (Liliaceae)

Quantitatively evaluated C-banded and conventional karyograms are presented for the related speciesScilla bifolia subsp.danubialis Speta,S. drunensis (Speta)Speta, andS. vindobonensis Speta. On the basis of banding patterns and karyotype structureS. bifolia subsp.danubialis (2n = 18, 2×) andS. drunensis (2n = 36, 4×) are quite similar, whileS. vindobonensis (2n = 18, 2×) is entirely different. There is a moderate degree of karyotypic variation withinS. bifolia subsp.danubialis andS. drunensis. However, inS. vindobonensis karyotypes and banding patterns are almost completely stable over a geographical range of about 500 km. The present results confirm the recent taxonomic separation ofS. vindobonensis fromS. bifolia, and suggest a considerable phylogenetic distance between these two diploid species. The results are discussed with reference to the morphological characters of the species and their geographical distribution.Evolution ofScilla and Related Genera, II.Dedicated to Univ.-Prof. Dr.Elisabeth Tschermak-Woess on the occasion of her 60th birthday.     

Origins of the new allopolyploid species Senecio cambrensis (asteraceae) and its relationship to the canary islands endemic Senecio teneriffae

The distribution of a 330 bp cpDNA insertion was reexamined in British material of Senecio cambrensis (2n = 6x = 60), its two putative parental taxa, S. squalidus (2n = 2x = 20) and S. vulgaris (2n = 4x = 40), and the closely related Canary Islands' endemic S. teneriffae (2n = 6x = 60). This formed part of a test of the hypothesis that the Welsh form of S. cambrensis is derived from introduced S. teneriffae rather than having originated in Wales via allopolyploidy as previously supposed. It was established that the 330 bp insertion was carried by all plants of Welsh S. cambrensis and also S. teneriffae, but was absent from the cpDNA of Edinburgh S. cambrensis and all plants of British S. squalidus and S. vulgaris var. vulgaris surveyed. However, two of 19 individuals tested of S. vulgaris var. hibernicus also possessed the cpDNA insertion, indicating that it is present in British material of S. vulgaris, although at low frequency. The close similarity between S. teneriffae and S. cambrensis, especially the Welsh form of S. cambrensis, was confirmed by the results of a restriction analysis of rDNA, and also morphometric and crossing studies. However, isozyme analysis showed that S. teneriffae is monomorphic for βEST-3 and ACO-1 phenotypes that are not present in Welsh and Edinburgh S. cambrensis, nor in S. squalidus and S. vulgaris material surveyed. It is concluded that S. teneriffae and S. cambrensis are two closely related allohexaploid taxa that have very similar, but different origins. It is postulated that whereas S. cambrensis (in Wales and Edinburgh) is the allohexaploid of S. vulgaris and S. squalidus, S. teneriffae is possibly the allohexaploid of S. vulgaris and S. glaucus, the latter being a diploid species closely related to S. squalidus.     

Chloroplast DNA as an evolutionary marker in thePhaseolus vulgaris complex

We have analyzed the changes occurring in the chloroplast DNA (cpDNA) of taxa belonging to thePhaseolus vulgaris complex to help clarify relationships among species of this complex. Two restriction maps for 11 restriction enzymes comprising the whole chloroplast genome from a wildP. vulgaris and a wildP. coccineus accession were constructed. These maps allowed us to compare a total of 330 restriction sites between the two genomes in order to identify polymorphisms, assess the type of mutations detected, and identify regions of high variability. A region, located in the large single-copy region near the borders with the inverted repeats, accounted for a large portion of the variation. Most of the mutations detected were due to restriction sites gains or losses. Variable and conserved regions were then evaluated in 30 accessions belonging to taxa of theP. vulgaris complex. Phylogenetic analyses were made using parsimony methods. Conclusions obtained from such analyses were the following: (1) there was high cpDNA variability withinP. coccineus but not inP. vulgaris. (2)P. coccineus subsp.glabellus showed a very distinct cpDNA type that strongly suggests that it actually belongs to a different but as yet undetermined section of the genus. Our cpDNA observations are supported by distinctive morphological traits and reproductive biology of this taxon. (3) InP. coccineus subsp.darwinianus (also classified asP. polyanthus), the cpDNA lineage was in disagreement with data obtained from nuclear markers and suggested a reticulated origin by hybridization betweenP. coccineus as the male parent and an ancestralP. polyanthus type, closely allied toP. vulgaris, as the seed parent. This initial cross was presumably followed by repeated backcrossing toP. coccineus. Our cpDNA studies illustrate the importance of molecular markers in elucidating phylogenetic relationships. They also indicate that accurate phylogenies will require analyses of both nuclear and cytoplasmic genomes.     

Even more new taxa from South and East Anatolia II

10 new Turkish taxa are described:Arenaria eliasiana, A. sivasica, A. monscragus, A. angustifolioides; Campanula lycica; Scutellaria orientalis subsp.tortumensis; Stachys choruhensis, S. tundjeliensis; Calamintha caroli-henricana; Aristolochia rechingeriana, the latter two species named in honour ofKarl Heinz Rechinger;Allium vuralii. Dedicated to Prof. DrKarl Heinz Rechinger on the occasion of his 80th birthday. For part I see Pl. Syst. Evol.154, 111–128.     

A numerical taxonomic study of thePolygonum aviculare complex (Polygonaceae) in Belgium

16 morphological characters were assessed in 300 plants sampled from 27 populations ofPolygonum aviculare sensu lato representative of the whole ecological range of the species in Belgium. The three multivariate treatments performed, namely principal component analysis, cluster analysis and discriminant analysis provide evidence thatP. aviculare can be divided in 4 units, roughly corresponding to the four taxa recognized byChrtek. However, the results point out that subsp.monspeliense and subsp.calcatum should be included at the varietal rank within respectively subsp.aviculare (=P. heterophyllum) and subsp.aequale (=P. arenastrum) as defined byLindman. Fruit dimensions, fruit shape and tepal length are the most discriminant characters for separating the four taxa, while several other characters are discriminant at the population level only. The evolutionary significance of the variation pattern of the whole complex is discussed in terms of life history differentiation and ecological specialization.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.     

Allozyme variability and phylogenetic relationships in the cultivated potato (Solanum tuberosum) and related species

Gene frequencies at 13 isozyme loci were determined in three South American taxa of cultivated potatoes [the diploid group (gp.) Stenotomum, the diploid subgroups (subgp.) Goniocalyx, and the tetraploid gp. Andigena ofS. tuberosum], in the diploid weed speciesS. sparsipilum, and in most of the main cultivars now raised in the Northern Hemisphere (the tetraploid gp. Tuberosum ofS. tuberosum). High levels of genetic variability (mean number of alleles per locus, percentage of polymorphic loci, and mean heterozygosity) were detected, being higher in tetraploid potatoes. An equilibrium among the evolutionary factors which increase genetic variability and artificial selection for maximum yield would explain the high uniformity of heterozygosity values we observed in both Andigena (0.36 ± 0.02) and Tuberosum (0.38 ± 0.01) cultivars.—The low value of genetic distance (D = 0.044) between Stenotomum and Goniocalyx does not support the status of species forS. goniocalyx.—In most isozyme loci, the electromorphs of gp. Andigena were a combination of those found in both gp. Stenotomum andS. sparsipilum, suggesting an amphidiploid origin of gp. Andigena from that two diploid taxa. The presence in Andigena of unique electromorphs, which were lacking in both gp. Stenotomum andS. sparsipilum, suggests that other diploid species could be also implied in the origin of tetraploid Andean potatoes. Furthermore, since Andigena were more related to Stenotomum (D = 0.052) than toS. sparsipilum (D = 0.241), the autopolyploidization of Stenotomum individuals and the subsequent hybridization with gp. Andigena may also have occurred. Thus, our study suggests a multiple origin (amphidiploidy, autoploidy, and hybridization at tetraploid level) of gp. Andigena.—Most of the electromorphs of gp. Tuberosum were also found in gp. Andigena; both the direct derivation of that group from the Andean tetraploid potatoes and the repeated introgression provided by breeding programmes could explain this result. However, the allele c of Pgm-B, present in 30 out of 76 Tuberosum cultivars from Northern Hemisphere as well as in 3 Chilean Tuberosum cultivars, lacks in the 258 Andigena genotypes sampled, suggesting that Chilean germplasm could have taken part in the origin of at least the 39% of the potato cultivars from Europe and North America analyzed here.—The distanceWagner procedure provides an estimate of a 30% of heterogeneity in the evolutionary divergence shown by different groups of cultivated potatoes. Diploid groups show a higher (22.5%) evolutionary rate than tetraploids, which can be attributed to both tetrasomic inheritance and facultative autofecundation that exists in Andigena and Tuberosum groups. Thus, artificial selection acting since 10000 years has not resulted in a higher rate of molecular evolution at the isozyme level in the tetraploids.     

On subspecific taxa ofHypericum empetrifolium (Hypericaceae) from Crete

The morphology, altitudinal distribution and phytocoenology ofHypericum empetrifolium Willd., var.oliganthum Rech. fil. and var.tortuosum Rech. fil. are reassessed and as a result, the taxa are treated at subspecific rank. Essential characters proved to be growth form, number of flowers, foliage and epidermal surface characters, shape of seeds and testa sculpturing.The different growth forms can be interpreted as adaptations to the local habitats.Hypericum empetrifolium Willd. subsp.empetrifolium is a low subshrub of various vegetation types known as phrygana whereasHypericum empetrifolium Willd. subsp.oliganthum (Rech. f.)Hagemann grows as a cushion-like dwarf shrub in chasmophyte associations rich in relic species.Hypericum empetrifolium Willd. subsp.tortuosum (Rech. f.)Hagemann is confined in the high mountain regions and grows as a true dwarf espalier shrub with low spreading shoots.Dedicated to Hofrat Univ.-Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

Delimitation and classification ofBromus fasciculatus (Poaceae)

The syntypes ofBromus madritensis var.delilei Boiss. comprise two different elements: specimens ofB. haussknechtii Boiss. and ofB. fasciculatus C. Presl s.l. By its lectotypification and on the basis of new morphological characters, the validation ofB. fasciculatus subsp.delilei (Boiss.)H. Scholz 1971 was substantiated. This subspecies represents the eastern marginal segregate of the south-Mediterranean species.B. fasciculatus var.alexandrinus Thell. is a minor variant of the typical subspecies.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

祁连山区特有物种黑虎耳草的居群遗传多样性研究

利用叶绿体基因(rbcL和trnS-G)及核糖体DNA内转录间隔区(ITS),对祁连山区狭域分布的特有物种黑虎耳草8个居群(115个个体)进行遗传多样性研究,旨在揭示黑虎耳草的居群遗传结构及其历史进化过程。结果表明:(1)所有取样个体共检测到4个cpDNA单倍型和9个ITS单倍型,其中祁连山东南部的居群固定较多的单倍型和特有单倍型,而西北部居群只固定少数几个广泛分布的单倍型,且遗传多样性普遍较低。(2)基于cpDNA数据和ITS数据的分子变异分析(AMOVA)显示,遗传变异主要来源于居群内。(3)基于cpDNA数据的中性检验表明,Tajima’s D(-1.012 30,P> 0.05)和Fu&Li’s D*(-2.066 77,P> 0.05)均为负值,均不显著;歧点分布分析结果显示,黑虎耳草居群经历过近期的扩张事件。根据物种现有遗传分布格局推测,黑虎耳草在第四纪冰期时退缩到祁连山东南部的边缘避难所,间冰期或冰期后回迁到祁连山西北部地区,在回迁过程中由于奠基者效应导致祁连山西北部的居群仅固定少数广泛分布的单倍型,并呈现出较低的遗传多样性;由于居群较小且相互隔离,该...     

Stachys beckeana (Labiatae) in Albanien und Jugoslawien

Morphological and cytological investigations were carried out inStachys beckeana Dörfler & Hayek andS. recta L. subsp.sarajevensis sensuHayek. These two taxa have the same morphological characteristics (shape and indumentum of leaves; morphology of sepals, petals and fruits) and both have 2n = 2x = 34 chromosomes. They cannot be considered as different taxa;S. beckeana Dörfler & Hayek is the valid name (synonyms:S. recta L. subsp.sarajevensis sensuHayek,S. hayekii Malý,S. recta L. subsp.hayekii Malý).

Erster Beitrag zur Neubearbeitung der Artengruppe derStachys recta L.