Nutlet sculpturing of selectedTeucrium species (Lamiaceae): A character of taxonomic significance

Nutlet sculpturing of 23Teucrium species belonging to six sections was studied. The nutlets of analysed species were characterized by the presence of the oil glands of various density. SectionTeucrium is well defined by very densely distributed trichomes and oil glands on nutlets. Trichomes on nutlets were absent in all other species, except inT. marum from sect.Chamaedrys. Sect.Polium is a homogeneous group, distinguished from the other sections. There are some similarities of this sect. with sect.Scordium. Species of sect.Chamaedrys are heterogeneous and overlap according to the surface sculpturing characters with the other sections. The significance of nutlet microcharacters as additional taxonomical markers in the infrageneric classification of the genusTeucrium is discussed.     

Mikromorphologie der Samenoberflächen der GattungEuphorbia

Seed coat surfaces of 127 species, representing 23 sections of the genusEuphorbia L. (Euphorbiaceae) have been examined with the scanning electron microscope (SEM). Various surface patterns and cell wall structures are described. In some groups the seed coat is rather uniform (e.g. in sect.Euphorbium), in other sections even closely related species can be separated by seed coat morphology (e.g.Tithymalus). The taxonomic application of testa micromorphology and the possible systematic position of certain taxa are briefly discussed. The seed coat cells of all species in sect.Tithymalus show remarkable intercellulares filled with small particles, which are described in detail for the first time.

     

Molecular phylogenetics ofCochlearia (Brassicaceae) and allied genera based on nuclear ribosomal ITS DNA sequence analysis contradict traditional concepts of their evolutionary relationship

The systematics and phylogeny of the genusCochlearia and allied genera are unsettled. There are no clearly defined genera and subtribal structures to determine subtribeCochleariinae in respect to subtribeThlaspidinae. The use of morphological data, such as fruit form or embryo characters have resulted in contradictory taxonomic concepts in the past due to their homoplastic nature. We investigated all sections of genusCochlearia recognised in the most common concepts, as well as some genera such asIonopsidium, Bivonaea, Pastorea andThlaspi s. l. pro parte. Previous studies based on molecular data and morphological studies have shown close relationships between taxa from subtribeCochleariinae andThlaspidinae. The Internal Transcribed Spacer regions of the nuclear encoded ribosomal DNA operon and the plastidictrnL intron were sequenced from a number of genera. A molecular phylogeny was derived and compared to traditional classification systems. These data grouped sections ofCochlearia outside theCochlearial Ionopsidium core group and integrated them either closely to genusNoccaea in subtribeThlaspidinae (sect.Pseudosempervivum) or positioned them outside both theCochlearia core group and theThlaspi s. l. clade (sect.Hilliella). The molecular data indicate that subtribal arrangements in tribeLepidieae are artificial and do not reflect evolutionary history. The genusCochlearia is represented by sectionsCochlearia andGlaucocochlearia and the genusIonopsidium should be integrated intoCochlearia.     

Nutlet morphology in Hemigenia R.Br. and Microcorys R.Br. (Lamiaceae)

Nutlets of Hemigenia R.Br. and Microcorys R.Br. were examined using SEM. Significant variation, mainly useful at the infrageneric level, was found in nutlet shape, nature of the attachment scar, nature of surface sculpturing, exocarp cell shape and sculpturing, and nature of the indumentum. Typical nutlets are ovoidal, strongly reticulate or rugose. The exocarp cells are isodiametric and convex to papillate. Also common are cylindrical nutlets, often with longitudinal ridging and papillate exocarp cells. Surface pitting and concave exocarp cells are rare. A cladistic analysis of nutlet characters suggests both Hemigenia and Microcorys are polyphyletic, and Microcorys paraphyletic with respect to Westringia Sm. Notwithstanding that, the infrageneric classification of Hemigenia was largely supported, while in Microcorys, there was support for sect. Hemigenioides, but sects Anisandra and Microcorys were not resolved as distinct.     

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.     

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.     

Analysis of the fruit surfaces in Bupleurum L. (Umbelliferae) with SEM

20 Bupleurum L. taxa were investigated for micromorphological typology of the fruit surfaces with SEM. Considerably different types of sculpturing at species and subspecies level and similarities between related taxa were observed. A dichotomic key was prepared using the diagnostic surface patterns. Reticulate (sect. Bupleurum), rugose (sect. Aristata Godron subsect. Aristata) and rugulose-colliculate-verrucate (subsect. Juncea Briq.) types of surface structures were distinguished. Micromorphological characteristics may be valuable additional taxonomical criteria for the delimitation of infrageneric groups of species.     

A contribution to taxonomy of Turkish <Emphasis Type="Italic">Linum</Emphasis> based on seed surface patterns

Seed samples of eighteen taxa from four sections in Linum were examined for diagnostic and taxonomic utility of the surface microstructures. Considerably different patterns were distinguished at specific and infraspecific levels. Similar sculpturings in closely related taxa and some common surface features for sectional groupings were observed. Smooth surface textures in sect. Linastrum, granulate-ruminate in sect. Dasylinum and rugulose structure in sect. Linum were typical characteristics. Sect. Syllinum exhibited miscellaneous patterns. Seed coat sculptures may be utilized as additional consistent parameters in the infrageneric delimitations and taxonomical association of Linum.     

Molecular systematics of the Old World Astragalus (Fabaceae) as inferred from nrDNA ITS sequence data

This study represents a nuclear rDNA ITS-based phylogenetic analyses of a greater sampling of the Old WorldAstragalus compared to our previous work (212 vs. 134 taxa). Phylogenetic relationships among 212 species (213 accessions) of the Old WorldAstragalus, including newly segregated monotypic genusPodlechiella, the two aneuploid New WorldAstragalus, and five related genera, were inferred from analyses of nuclear rDNA ITS sequences using maximum parsimony. A total of 658 nucleotide sites and four binary characters for indels were analyzed. The results of phylogenetic analyses suggest sect.Phyllolobium, comprising mostly the Chinese species, is placed outside of the so-calledAstragalus s. str. and is a well-supported monophyletic group. The monotypic annual segregate genusThlaspidium (≡Astragalus sect.Thlaspidium, A. thlaspi), is clearly nested withinAstragalus s. str. Among the many sections analyzed here, only sects.Cenanthrum, Caraganella, Eremophysa, Incani, Laxiflori, andLotidium are strongly supported as monophyletic. Our analysis, in agreement with previous studies, shows that the North American euploidAstragalus species are scattered throughout the Old World groups of the genus.     

Phylogenetic relationships in theSideritis leucantha group (Lamiaceae)

Six closely related taxa of the sect.Eusideritis of the genusSideritis (S. leucantha, S. pusilla, S. flavovirens, S. granatensis, S. biflora andS. osteoxylla) are analysed to elucidate their phylogenetic relationships and position within the sect.Eusideritis. Meiotic behaviour, karyotype features, size and fertility of pollen grains, DNA amounts and seed protein profiles are reviewed. A polyploid origin of the group (from x = 7) and the further diversification through dysploidy and chromosome repatterning is postulated.S. osteoxylla is apparently of hybrid origin.     

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.     

The pollen morphology ofOdontites (Scrophulariaceae) and its taxonomic significance

The pollen morphology of 29 species ofOdontites and related genera was examined by light and scanning electron microscopy. Three major pollen types differing fundamentally in exine sculpturing were found. In the parasitic tribePedicularieae retipilate sculpturing is the most widespread type representing a plesiomorphic character state from which the two other types are derived. Of these, reticulate sculpturing is confined toOdontites, whereas a complex retirugulate pattern was found only in the monotypic Near East genusBornmuellerantha. Within the retipilate and the reticulate major types eight minor types were distinguished based on the differential correlation of exine surface morphology, size, shape and amb form. The pollen data are generally well correlated with macromorphological features and furnish important taxonomic characters at the genus—as well as at the specieslevel.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCanina G. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.     

Phylogeny ofRubiaceae-Rubieae inferred from the sequence of a cpDNA intergene region

A phylogenetic analysis of 25 species, representing eight genera of theRubieae tribe (Rubiaceae), has been made using the DNA sequence of the chloroplastatp B-rbc L intergene region. Six tropical genera from other tribes ofRubiaceae have been used as outgroups. Whatever the method of analysis (distance, parsimony or maximum likelihood), five groups are clearly separated and described as informal clades. Their relative relationships are not clearly resolved by the parsimony analysis, resulting in eight equally parsimonious trees, 327 steps long, with a consistency index (CI) of 0.749 (excluding uninformative sites). TheRubieae tribe appears monophyletic from the data available. Some new and partly unexpected phylogenetic relationships are suggested. The genusRubia forms a separate clade and appears to be the relatively advanced sister group of the remaining taxa. TheSherardia clade also includes the generaCrucianella andPhuopsis. Galium sect.Aparinoides appears closely attached to theAsperula sect.Glabella clade. The remaining taxa ofGalium are paraphyletic:Galium sect.Platygalium (in theCruciata clade) is linked to the advanced generaCruciata andValantia; the more apomorphic groups ofGalium form theGalium sect.Galium clade, including the perennial sectionsGalium, Leiogalium, andLeptogalium as well as the annual (and possibly polyphyletic) sect.Kolgyda.     

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.     

Enzyme analysis of genetic variation and relationships in diploid and polyploid taxa ofGalium (Rubiaceae)

Allozyme variation at 11 loci (with 37 alleles) was studied electrophoretically in seven outbreeding, closely related diploid and tetraploid taxa, seven from sect.Leptogalium and two from sect.Leiogalium. Whereas the sections are clearly distinct by several different alleles, aggregates, species and subspecies differ only in the frequency or presence/absence of common alleles. The resulting dendrogram suggests phylogenetic relationships and is supported by other multidisciplinary evidence. Tetraploids have originated independently in several groups, and there is evidence for tetrasomic inheritance and thus for autopolyploidy in spite of normal meiotic bivalent pairing and partly suspected hybrid origin. Tetraploids differ from related diploids only little in number of alleles and expected heterozygosity within populations, but clearly exhibit higher numbers of genotypes. This often corresponds to their greater morphological variability, increased adaptive flexibility, and better colonizing capacity compared to related diploids.     

Pollen morphology of Stachys (Lamiaceae) in Iran and its systematic implication

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.