Relationships between seven species ofVulpia (Poaceae)

Seven species ofVulpia were subjected to morphological and protein studies using principal component and cluster analysis and theJaccard's Similarity Coefficient. Studies showed all species were uniform and distinct from each other. Species of sect.Loretia were very dissimilar from those of sect.Monachne and sect.Vulpia. The latter section comprises taxa of high mutual affinity.     

Species relationships inFestuca (sect.Ovinae,Poaceae)

Morphological and seed protein studies of selected species ofFestuca were performed to elucidate the relationships between species of sect.Ovinae and their affinity with other fescues.Festuca rubra andF. heterophylla (extravaginale group) had higher phenetic affinity with taxa of sect.Scariosae, Montanae andBovinae than with members of their own section. 5 species of the intravaginale group (F. ovina, F. capillata, F. valesiaca, F. sulcata, F. trachyphylla) were fairly uniform and well defined by protein and morphological data. This supports current systematic thinking and does not agree withHackel's (1882) classification of this group.     

A reexamination of infraspecific taxa of a wild potato,Solanum microdontum (Solanum sect.Petota:Solanaceae)

Current taxonomic interpretations ofSolanum microdontum Bitter partition the species into two or three infraspecific taxa, variously recognized as subspecies or varieties. The present study reexamines these taxa using morphological data from four individuals each of 69 accessions from most of the range of the species, planted in a common field plot. Our results show that the character states used to recognize infraspecific taxa inS. microdontum often vary within accessions and have no correlation with geography. We conclude that past hypotheses have used typological concepts and that infraspecific taxa are not warranted. This study questions other hypotheses of infraspecific taxa in sect.Petota.     

Ecogeographical differentiation patterns inAdonis sect.Consiligo (Ranunculaceae)

Adonis sect.Consiligo is split into three subsections based on morphology, growth pattern, pollen and fruit morphology. For all species distribution maps are provided and discussed with regard to their ecology and climatic requirements. These data reveal additional characters that support the classification of the taxa. Most species of the section grow in continental Eurasiatic steppes and mountain steppe habitats. Only the species of subsect.Amurenses grow in Far Eastern temperate forests and prefer a more oceanic climate. The taxonomic position of these strongly divergent species is supported by the distributional and ecological data. The evolution ofAdonis sect.Consiligo occurred under strong climatic control. During evolution the taxa could only slightly widen their ecological amplitude and, thus, ecology remained rather stable except that of the subsect.Amurenses. Ecological and distributional data can be used to validate taxonomy and phylogeny and reveal additional informations.     

Phenetic investigation of non-nodulating African species ofAcacia (Leguminosae) using morphological and molecular markers

Morphological and RAPD markers were used to assess the relationships among nodulating and non-nodulating species of AfricanAcacia. Non-nodulating species of AfricanAcacia are only found within subg.Aculeiferum sect.Monacanthea. African species of sect.Monacanthea examined were found to form a group distinct from the other African species examined on a morphological and molecular basis. All lack the ability to nodulate, suggesting that non-nodulation may be used as a taxonomic tool. The species of sect.Aculeiferum were separated by RAPD and morphological analysis into two groups depending on whether they were armed with prickles in pairs and/or prickles in threes, or solitary. A third group of species was identified within sect.Acacia: further subdivision of this group was achieved into subsectt.Pluriseriae andUniseriae. The position ofA. albida relative to other AfricanAcacia species was found to be distinct but not totally independent of the genus. The partitioning and distribution of the genetic variability within the genus is further elucidated by the RAPD analysis of populations ofAcacia species. A population analysis ofA. polyacantha demonstrated geographical and site-specific variation.     

Synopsis ofVerbesina sect.Ochractinia (Asteraceae)

A taxonomic revision ofVerbesina sect.Ochractinia is presented. 40 species plus 7 additional infraspecific taxa are recognized. A table of the distribution of the species, key to the species and synonymy are included. Relationships to other sections ofVerbesina and species relationships within sect.Ochractinia are discussed.     

Biflavonoid differentiation in sixBartramia species (Bartramiaceae,Musci)

The occurrence of eight biflavones belonging to the biluteolin series, the apigenin-luteolin series and the 2,3-dihydro-biluteolin series, and one monomeric flavone with an acid group, was investigated in six taxa ofBartramia with emphasis on sect.Ithyphyllae. The variation of biflavone profiles contributes to the characterization of theBartramia species studied; substitution patterns signalize relevance also on the sectional level. Based upon the flavonoid composition,Bartramia afro-ithyphylla is suggested to be transferred from sect.Ithyphyllae to sect.Bartramia of the genusBartramia.     

New variation and biosystematic patterns detected by allozyme and morphological comparisons inLimnanthes sect.Reflexae (Limnanthaceae)

Sixty-one populations ofLimnanthes sect.Reflexae were surveyed for variation at 19 allozyme loci and reassessed for their morphological diversity and biosystematic relationships. Cluster analysis of allozyme data at individual population level provided evidence for a fairly good fit with the earlier systematic work on this genus but both morphological and allozyme data together defined some new clusters as well as exceptional entities. For example, only var.rosea of the fourL. douglasii varieties described byMason (1952) on the basis of geographical distribution, flower color and leaf morphology, remained as a distinct entity; the others were realigned along new geographic boundaries.L. striata appeared to have two well-developed subgroups, one of which was closely related to theL. douglasii complex. Further ecogeographic and hybridization studies are needed to fully describe these new phenetic findings in the context of phylogenetic inference. SectionInflexae was readily distinguished from sect.Reflexae by several diagnostic loci, thus supporting the validity of this division. Interspecific variation levels and population differentiation inLimnanthes were high when compared to other plant taxa, possibly indicating highly dynamic evolutionary processes for which variable habitats and extreme subdivision within species have been suggested as explanations.     

Fruit differentiation, palynology, and systematics in theScabiosa group of genera andPseudoscabiosa (Dipsacaceae)

Fruits ofDipsacaceae are single-seeded, have bristle-shaped calyx segments and are tightly enclosed by four fused bracts forming an epicalyx. Comparative morphological and anatomical studies reveal a great diversity of epicalyx and calyx, often relevant to fruit dispersal. The present contribution deals with theScabiosa group of genera, the core of theScabioseae tribe. Most of its taxa develop a diaphragma from a meristem on the inside of the epicalyx. This diaphragma, together with the lower part of the epicalyx encloses the fruit proper, whereas the upper parts form a so-called epi-diaphragma (ed) and a ± hyaline corona. Differences of the epicalyx with respect to the size and position of the ed, elaboration of the corona, origin of pits (=foveoles) and other morphological and anatomical specializations can be demonstrated. Together with palynological and karyological data these new facts support an improved concept of relationships and systematics for the taxa studied:Scabiosa sect.Scabiosa and sect.Cyrtostemma are closely related and should be united to form the genusScabiosa s. str.;Pycnocomon can be maintained as an independent genus, sister toScabiosa sect.Trochocephalus which then has to be treated as a genus,Lomelosia. In contrast, the following genera have to be included inLomelosia:Tremastelma asLomelosia sect.Callistemma, andScabiosiopsis as part ofLomelosia sect.Lomelosia. Pseudoscabiosa deviates in so many features that it has to be excluded from the redefinedScabioseae s. str.     

Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

Utility of rDNA ITS sequences in the systematics ofTeucrium sectionPolium (Lamiaceae)

A phylogenetic study based on sequence data from the complete internal transcribed spacer region (ITS) of nuclear ribosomal DNA for sect.Polium of the genusTeucrium shows both intersectional congruence and intrasectional incongruence with traditional morphological classifications. We attribute this incongruence largely to problems related to the polyploid complex studied. SectionPolium includes many poorly differentiated taxa of probable recent origin through hybridization followed by polyploidization. Both on the basis of parsimony and distance (Neighbor-Joining method) analyses,T. dunense andT. thymifolium are the species that diverge most from the sampled taxa. However, unlikeT. thymifolium, the morphology ofT. dunense is not much differentiated in relation to the other taxa. Both species are, nonetheless, the only sampled species to occupy isolated, exclusive environments and which may have undergone rapid evolution by a bottleneck effect.Teucrium dunense is found only on dunes along the Spanish and French coasts andT. thymifolium, a chasmophytic species with limited endemism, is found solely on limestone and dolomite cliffs in the low mountains in south-eastern Spain. A hypothesis is presented to explain the discrepancy between the observed comparatively large amount of ITS sequence divergence and the low morphological differentiation inT. dunense.     

Molecular phylogeny ofIva (Asteraceae,Heliantheae) based on chloroplast DNA restriction site variation

Iva s.str. (comprising ten species) was examined by cpDNA restriction site variation to determine phyletic relationships within the group. The results were compared with relationships proposed from other data. A total of 86 restriction site mutations was detected, 47 of which proved phylogenetically informative. A single most parsimonious tree was obtained using both Wagner and Dollo parsimony. The tree revealed three main lineages that are congruent with the three chromosome lineages (base numbers of x = 16, 17, 18). The monophyly of the x = 16 and 18 groups was supported strongly by molecular data, while the monophyly of x = 17 lineage was only supported moderately. Relationships among the three lineages indicate that the sect.Iva is paraphyletic because sect.Linearbractea is nested within it. Both morphological data and the secondary chemical data are in agreement with the proposed cpDNA phylogeny. Because of this agreement, sect.Iva is revised such that,I. axillaris was excluded and positioned within the newly proposed sect.Rhizoma. Patterns and rates of cpDNA evolution were also examined. The results indicated an uneven evolution in the chloroplast genome with different rates of cpDNA evolution in at least a few species ofIva. However, the evolutionary clock hypothesis can not be rejected within most of the lineages inIva.     

Sero-systematics ofCytisus sect.Trianthocytisus (Fabaceae)

The serological reaction of seed proteins provides evidence for a partly new systematic arrangement ofCytisus sect.Trianthocytisus and ofCytisus s.l. Proposed modifications agree with recent advances in morphological taxonomy. Sect.Trianthocytisus includes only two species,C. villosus andC. aeolicus. Its position is central within the genus, and this fact agrees with the proposed retypification ofCytisus (type species:C. villosus).C. emeriflorus, formerly included in the same section, constitutes the monospecific sect.Emeroides, which is intermediate towards the genusLembotropis. This is serologically isolated and includes onlyL. nigricans. It is confirmed thatC. sessilifolius should be removed from the genusCytisus as a monospecific genus:Cytisophyllum Lang which is closely allied toHesperolaburnum and toPodocytisus, the most primitive genera ofGenisteae.     

The potential of AFLPs in biosystematics: A first application inSolanum taxonomy(Solanaceae)

Using the AFLP technique highly informative DNA fingerprints were generated from 19 taxa ofSolanum sect.Petota (potatoes) and three taxa ofSolanum sect.Lycopersicum (tomatoes). Both phenetic and cladistic analyses were conducted from the individual genotypic level to the species level. An AFLP fingerprint, using a combination of suitable AFLP primers, generated 12 to 71 scorable fragments per genotype which was sufficient for taxonomic interpretation. The classifications based on the molecular markers were generally in agreement with current taxonomic opinions. Unexpectedly,S. microdontum was associated with ser.Megistacroloba rather than with ser.Tuberosa, andS. demissum (ser.Demissa) and species of ser.Acaulia appeared closely affiliated. AFLP is an efficient and reliable technique to generate biosystematic data and therefore a promising tool for evolutionary studies.     

Electrophoretic seed albumin patterns inVicia species of sectt.Hypechusa andPeregrinae (Fabaceae)

This work is a continuation of electrophoretic investigations aimed at revealing a wild relative ofVicia faba. Electrophoretic analysis (PAGE) of seed albumins covered 52 accessions representing eightVicia species of sect.Hypechusa and two species of sect.Peregrinae. Most of the examined species showed an intraspecific variation due to differences between accessions and/or individual variation within accessions. In spite of the intraspecific variation, marked interspecific differences were recorded. However, none of the investigated species displayed electrophoretic seed albumin patterns similar to those reported earlier forV. faba. Contribution of the obtained results to characterization of the examined taxa is discussed.     

Systematic implications of chloroplast DNA variation inLepidium sectionsCardamon,Lepiocardamon andLepia (Brassicaceae)

Chloroplast DNA restriction site variation was examined amongLepidium taxa usually classified in sects.Lepia, Lepiocardamon andCardamon. By using 15 restriction endonucleases, filter hybridization experiments, and comparative mapping procedures, a total of 119 variable restriction sites was detected. Of these, 56 were phylogenetically informative and were used in a cladistic analysis. The resulting phylogenetic tree agrees with results derived from morphology, isozyme electrophoresis and the analysis of glucosinolates. The data suggest: 1) The recognition of sect.Lepia, and 2) that sect.Lepiocardamon has close relationships to sect.Cardamon and that both should be amalgamated. Low rates of interspecific chloroplast DNA sequence divergence in sect.Lepia correlated well with morphological data and isoelectric focusing analysis of Rubisco, suggesting that taxa of sect.Lepia are closely related and most likely have diversified recently.     

Phylogenetic analysis of the generaCladonia andCladina (Cladoniaceae, lichenizedAscomycota)

A cladistic analysis of 44 species of the generaCladina andCladonia is presented.Pycnothelia papillaria, Cladia aggregata andC. retipora were used as outgroup taxa. The consensus of all equally parsimonious trees suggests a common ancestral origin for species inCladonia andCladina while the generaPycnothelia andCladia cluster outside this group. The results do not support distinction ofCladina at genus level although it is distinguished as a monophyletic group but within the genusCladonia. The current sectional division ofCladonia is not supported. SectionsCocciferae andHelopodium are best supported but even these groups as currently delimited seem to in include elements that are more closely related to species of the other sections, e.g.C. carneola of sect.Cocciferae andC. pityrophylla of sect.Helopodium. Sections such asUnciales andPerviae seem to be artificial assemblages. In general the evolutionary scheme of the genus seems to be more complicated than revealed by the current sectional division.     

RAPD patterns,nrDNA ITS sequences and morphological patterns inSilene sectionSedoideae (Caryophyllaceae)

Hierarchical patterns inSilene sect.Sedoideae were investigated using random amplified polymorphic DNA (RAPD), nucleotide sequences of the internal transcribed spacer (ITS) regions of the 18S–28S nuclear ribosomal DNA, and discrete morphological characters. All data sets firmly supported the species recognized. The RAPD data offered the best resolution at the intraspecific level, supporting the current intraspecific classifications ofS. sedoides andS. integripetala. The ITS sequences and the morphological data gave poor resolution within species, and the three data sets disagreed about the relationships among species. The signal from the RAPD data was strongest and remained when the total data set was analysed. The three data sets all support an amphiploid origin ofS. aegaea, with the strongest evidence from the ITS sequences. Incongruences among data sets as well as merits and shortcomings of each are discussed. The robustness of the results can be evaluated using perturbations of data, i.e., bootstrap and jackknife of taxa and characters. These methods should not be taken as methods of statistical inference at the taxonomic level, because unbiased sampling appears impossible. RAPD data, however, come close to being suitable for statistical estimation of hierarchies at the genome level, but several methodological problems have to be solved.     

Hybridisation,genomic constitution and generic delimitation inElymus s. l. (Poaceae: Triticeae)

Intergeneric crosses were made between representatives of the genomically-defined generaElymus, Agropyron, Elytrigia, Pseudoroegneria, andThinopyrum. The genomic constitution ofElytrigia repens, the type species ofElytrigia, is shown to be SSH, a genomic combination otherwise found only inElymus. The S genome ofPseudoroegneria has almost always a dominant influence on the morphology of the taxa of which it is a component.Wang (1989) showed that the J genome inThinopyrum and the S genome have considerable homoeology, with a mean c-value of 0.35 in diploid SJ hybrids. A genetic coherence from S to SJ^e, J^e, J^eJ^b, and J^b can be expected, agreeing with the continuous morphologic variation pattern observed. Because of the absence of morphological discontinuities between the taxa,Pseudoroegneria (S),Elymus (SH, SY, sometimes with additional genomes),Elytrigia (SSH, SSHX), andThinopyrum (SJ, SJJ, J) are best treated as a single genus,Elymus, following the generic concept ofMelderis in Flora Europaea and Flora of Turkey. The basic genomic constituents ofElymus will then be the S and/or J genomes.Agropyron, with diploids, tetraploids, and hexaploids based on the P genome is morphologically distinct from other genera inTriticeae. In a few species ofElymus andPseudoroegneria, a P genome is an additional constituent. In these cases the P genome has a negligible morphological influence. Therefore, it seems reasonable to maintainAgropyron as a separate genus.     

Variation in wild populations of annual beet (Beta,Chenopodiaceae)

Among the morphological variation in wild annual populations of sect.Beta only tepal characters show a geographic pattern, and hence can be used to distinguish different taxa. Two morphological types correspond with taxa already described:B. macrocarpa (incl.B. bourgaei) has a Macaronesian and W. to E. Mediterranean andB. adanensis an E. Mediterranean distribution area. A third type in the Aegean region is not well known yet and possibly has to be included inB. macrocarpa. Both diploid and tetraploid (x = 9) cytotypes are found withinB. macrocarpa, the latter exclusively on the Canary Islands.