Differentiation patterns in the E. MediterraneanScutellaria rubicunda group (Lamiaceae)

Strict sterility barriers where found between theScutellaria populations on Sicily and the Greek populations. The material from Samos is separated by a strict sterility barrier from the other Greek populations. In contrast, the populations on mainland Greece and adjacent islands are all ± interfertile. Three species are recognized,S. rubicunda Hornem., endemic to Sicily, andS. brevibracteata subsp.icarica, endemic to Samos and Ikaria. All other populations are referred toS. rupestris with eight subspecies, most of which are endemic to one island or to one mountain. Three new subspecies are described, viz. subsp.rechingeri and subsp.olympica, endemic to mt Vourinos and mt Olympus in North Greece, respectively, and subsp.caroli-henrici, native to the Malea peninsula of Peloponnisos. The phytogeographical connections and genetic differentiation within and between populations are discussed.Dedicated to Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

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.     

Genetic relationships between European and Siberian larch,Larix spp. (Pinaceae), studied by allozymes. Is the Polish larch a hybrid between these two species?

Thirteen populations ofLarix decidua subsp.decidua and subsp.polonica, and three populations ofL. sibirica were investigated by starch-gel electrophoresis. In the populations assayed 61 alleles at 17 loci were revealed. The allozyme data support the earlier observations about close relationships between these two larch species. Nei's genetic distances betweenL. decidua andL. sibirica were relatively small (D = 0.057), however, almost five times larger, on average, than those between populations of the same species. Results obtained in this study disagree withBobrov's hypothesis about the hybrid origin of the Polish larch and suggest a direct origin from the European larch.     

A taxonomical revision ofThymus sect.Teucrioides (Lamiaceae)

TheThymus teucrioides Boiss. & Spruner aggregate is revised and the following new taxa, all from the alpine zone in the Greek mountains, are described:Th. leucospermus Hartvig from the calcareous mountains of Pindhos and Mt Parnassos in Sterea Ellas,Th. rechingeri Hartvig with the subsp.macrocalyx Hartvig from calcareous mountains in Sterea Ellas and N Peloponnissos, andTh. teucrioides subsp.alpinus Hartvig from the serpentine areas of N Pindhos. In the variableTh. teucrioides s. str. many characters have turned out to be markedly geographically correlated and many local populations can be distinguished by a particular combination of characters.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

Taxonomy and distribution of Poa carpatica in the Western Carpathians

We examined the taxonomic status and distribution of Poa populations from supramontane and subalpine belt of the central Western Carpathians, so far classified as P. nemoralis subsp. carpatica and P. nemoralis subsp. montana. Significant morphological differences from P. nemoralis s. str. as well as combination of shared vs distinct characters allow us to attribute the populations under study to the species P. carpatica (V. Jirásek) Chopyk with two subspecies: P. carpatica subsp. carpatica a P. carpatica subsp. supramontana subsp. nova.     

Genetic introgression between tetraploidDactylis glomerata subspp.reichenbachii andglomerata in the French Alps. Insight from morphological and allozyme variation

The morphology, phenology and allozyme polymorphism in seven tetraploidDactylis glomerata populations growing in the French Alps were studied. Based on habitat and morphological characteristics, two of the populations could be classified asD. glomerata subsp.reichenbachii, which is known to comprise diploid and tetraploid plants growing exclusively on dolomite, on south-facing meadows. This subspecies has been previously recorded in the Italian and Swiss Alps but not in the French Alps. The remaining five populations were morphologically intermediate but more similar to the cosmopolitanD. glomerata subsp.glomerata. On the basis of allozyme variation, all populations were more related to subsp.reichenbachii than to the cosmopolitan subsp.glomerata, suggesting that gene flow has occurred between the two tetraploid subspecies. The possibility that selection has acted differently on morphological and allozyme characters is discussed.     

Chloroplast DNA variation of Panax (Araliaceae) in Nepal and its taxonomic implications

The restriction site and size variation of five PCR amplified fragments of noncoding chloroplast DNA (cpDNA) was examined in material from 13 populations ofPanax from Nepal and China. Fourteen restriction endonucleases produced 81 restriction site and length variations from the large single-copy region of cpDNA, 27 of which are polymorphic. The cpDNA dataset suggests two distinct groups ofPanax from Nepal (clades I and II). Clade I consists of two populations ofP. pseudoginseng subsp.pseudoginseng, and clade II is composed of material referable toP. pseudogingeng subsp.himalaicus (vars.himalaicus, angustifolius, andbipinnatifidus). The three accessions ofP. pseudoginseng subsp.japonicus andP. ginseng studied from China had cpDNA characters that differed from the HimalayanPanax. The highly distinctive cpDNA profile and morphology ofP. pseudoginseng subsp.pseudoginseng sensu Hara (1970) from central Nepal support its status as a separate species, which has an extremely restricted distribution.     

Chemical variation in two species of theLecanora subfusca group (Lecanoraceae,lichenizedAscomycotina)

The chemical variation inLecanora epibryon andL. subimmersa, two species of theL. subfusca group, has been examined. In both species the chemical differences are correlated with geographical distribution, but not with morphological differences. As a consequence the chemotypes are recognized at subspecific level. InL. epibryon three chemical races are segregated according to the various chemosyndromes present. Subspeciesepibryon, containing atranorin and triterpenoids, occurs in the northern hemisphere and South America, while the other two subspecies occur only in the southern hemisphere. The subsp.broccha (Nyl.)Lumbsch, comb. nova, contains atranorin, the stictic acid and the 2,5,7-trichloro-3-O-methylnorlichexanthone chemosyndromes, while subsp.xanthophora Lumbsch, subsp. nova, is similar but lacks the stictic acid chemosyndrome. Two chemical races occur in the pantropical speciesL. subimmersa. While subsp.subimmersa contains atranorin and zeorin, subsp.ramboldii Lumbsch & Elix, subsp. nova, contains an additional ten chlorinated xanthones.L. impressa is reduced to synonymy toL. subimmersa.     

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.     

Change of adaptations from entomophily to autogamy inParahebe linifolia (Scrophulariaceae)

Measurements of 16 inflorescence and floral characters from 5 populations ofParahebe linifolia (3 of entomophilous subsp.linifolia; 2 of autogamous subsp.brevistylis) were analysed. Discontinuous variation of three characters associated with the mechanisms of pollination suggests these are under strong selection towards different optima in each subspecies. Variation which is continuous and associated with the type of pollination in 4 characters mostly concerned with display is thought to result from selection for metabolic economies, counteracted strongly in subsp.linifolia, but less so in subsp.brevistylis, by selection for display. Nine more characters vary in a way not known to be associated with the two pollination systems.     

Taxonomic and morphological studies inHypecoum sect.Hypecoum (Papaveraceae)

Hypecoum sect.Hypecoum comprises 8 species, one with 2 additional subspecies, in the Mediterranean area and the Middle East. Five of these are recognized for the first time, viz.Hypecoum procumbens L. subsp.fragrantissimum Å. E. Dahl,H. procumbens L. subsp.atropunctatum Å. E. Dahl,H. trullatum Å. E. Dahl,H. angustilobum Å. E. Dahl andH. torulosum Å. E. Dahl. H. dimidiatum Delile andH. pseudograndiflorum Petr., which have generally been included inH. imberbe Sm., are reestablished as distinct species. Morphology and variation patterns are described and discussed with special reference to mating systems. Self-incompatibility is dominating butH. procumbens subsp.procumbens andH. torulosum are self-compatible. The broad variation ranges of the former taxon in traits presumably related to mating system (petal, anther and stigma sizes as well as pollenovule ratios) indicate varying rates of outcrossing. SEM micrographs are given of petal and stigma surfaces and of pollen grains. Chromosome counts are presented for seven taxa. All are diploid with 2n = 16.     

Biometrical variability of foliage and cone characters inCupressus bakeri (Cupressaceae)

Biometrical variability of foliage, cones, and seeds were analyzed in 8 native populations (throughout the range) of a relictual species,Cupressus bakeri Jeps. (Cupressaceae) from California and Oregon. The partition of variation within and among populations for this rare plant was investigated and the justification for subspecies status was reviewed. Morphological characters varied significantly, with more of the variation among populations than at the population level. Multivariate analyses indicated close affinities among populations on serpentine sites suggesting the presence of a serpentine race. However, the high degree of variation among populations did not permit their clear separation into the two subspecies described byWolf (1948). We conclude thatC. bakeri should be treated as a monotypic species.     

Morphometrics and character state recognition for cladistic analyses in theBletia reflexa complex (Orchidaceae)

Different criteria have been suggested for decisions on character state recognition for morphological characters showing continuous variation from taxon to taxon. A method proposed byMishler andDe Luna seeks to recognize as many character states as the groups of taxon means that are revealed by an analysis of variance (ANOVA) followed by multiple range tests. We apply this method as a guide in decisions of selecting taxonomic characters in species of theBletia reflexa complex. Seven species of the complex, represented by 156 individuals belonging to 54 populations, were the terminal taxa of the study. Character state codes were assigned to 20 characters. These characters together with 12 non-quantitative characters were incorporated into cladistic analyses. Results indicate that the complex is monophyletic only under one outgroup combination.     

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.     

A revision of European taxa ofHyacinthoides sectionSomera (Hyacinthaceae) on the basis of multivariate analysis

Cluster analysis and principal components analysis were used to investigate patterns of morphological variation among specimens of European species of sect.Somera of the genusHyacinthoides (Hyacinthaceae). The results are clearly consistent with the existence of four species (H. italica, H. paivae, H. reverchonii andH. mauritanica). It is proposed that the latter be considered to comprise two subspecies,H. mauritanica subsp.mauritanica andH. mauritanica subsp.vincentina (comb. nov.). A key is presented, together with distribution maps for the five taxa.     

The vicarious differentiation of the alpineGalium serpylloides group (Rubiaceae), endemic to the W. Himalaya

Galium serpylloides agg. comprises a group of locally endemic, disjunct and vicarious taxa in the (sub)alpine zone of the W. Himalaya, from W to E:G. gymnopetalum, G. lahulense, G. serpylloides s. str.,G. saipalense, andG. nepalense. Four of these are new to science;G. serpylloides is newly typified. The main characters of the five species are compared and illustrated, their affinities and evolution are discussed, and a map of distribution is presented.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

Multivariate morphometric and allozymic analysis of theConospermum taxifolium (Proteaceae) species complex

A morphometric analysis of 18 attributes of 110 plants of theConospermum taxifolium complex suggests that it consists of three polythetically distinct taxa, corresponding to the traditional concepts ofC. taxifolium Smith s. str.,C. ericifolium Smith andC. ellipticum Smith. Discrimination is possible on the basis of leaf but not flower attributes. Analysis of allozymic variation indicates that the taxa are also genotypically differentiated.C. ericifolium andC. ellipticum are geographically isolated from each other but not fromC. taxifolium, andC. taxifolium is usually ecologically segregated from the other two taxa. Where this ecological segregation breaks down, morphological intermediates sometimes occur as the result of hybridization.     

Morphological and molecular comparisons ofCampsis grandiflora andC. radicans (Bignoniaceae), an eastern Asian and eastern North American vicariad species pair

Morphological and molecular comparisons were made forCampsis grandiflora (Thunb.)K. Schumann (Bignoniaceae) from eastern Asia andC. radicans (L.)Seemann from eastern North America. Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases. The cpDNA divergence between the two vicariad species was 2.44%, which is the highest reported among North Temperate disjunct taxa and one of the highest reported for infrageneric taxa. Detailed morphological comparisons also suggest a high level of divergence. Cluster analyses based on 22 morphological characters and 39 OTUs revealed two distinct groups corresponding with the two species. The average taxonomic distance between the two species was 1.806. Analyses of variance (ANOVA) revealed that 12 of the 18 quantitative characters differed significantly ( 0.01) betweenC. grandiflora andC. radicans. Divergence time based on cpDNA data was estimated as 24.4 million years. The Bering land bridge hypothesis was favored over the North Atlantic land bridge hypothesis based on the estimated divergence time and the geological history of the North Temperate region. The high levels of morphological and cpDNA divergence are not consistent with morphological stasis, which has been proposed as a common mode of evolution for North Temperate disjunct taxa.     

Phytochemical and morphological support for the existence of two species inMonoclea (Hepaticae)

Recognition of two different species in the liverwort genusMonoclea Hook. (monotypic orderMonocleales), viz.M. forsteri Hook. in New Zealand andM. gottschei Lindb. in the New World, is supported by characteristics of the sporophyte, antheridial receptacle and secondary metabolites.M. gottschei produces the greatest variety of flavonoids and the largest amount of bisbibenzyls ever encountered in a liverwort. In contrast,M. forsteri is poor in secondary metabolites. Two allopatric subspecies are recognized inM. gottschei, based on characteristics of the antheridial receptacle: subsp.gottschei in Chile (Valdivian region, Juan Fernandez Is.) and subsp.elongata Gradst. & Mues, subsp. nova, in tropical America. The exclusive occurrence inMonoclea of glucuronide and galacturonide flavone glycosides and the fact that capsule dehiscence may take place before full elongation of the seta are new arguments in support of the placement ofMonocleales in theMarchantiidae. Publication Nr. 43 of the Arbeitskreis Chemie und Biologie der Moose, Universität des Saarlandes, Saarbrücken. This paper is dedicated to DrElla O. Campbell, Massey University, Department of Botany and Zoology, New Zealand on the occasion of her 80th birthday.     

Biometrische Untersuchungen an Populationen vonOphrys cornuta,O. holosericea und ihren Hybriden (Orchidaceae)

Population variability ofOphrys holosericea (Burm. f.)Greut. subsp.holosericea (=O. fuciflora Crantz subsp.fuciflora) from near Vienna (Austria), and of subsp.maxima (Fleischm.)Greut. andO. cornuta Steven with intermediates from the Dalmatian island Hvar (Yugoslavia) was analysed and illustrated by scatter diagrams. A hybrid origin of these intermediates is suggested. Aspects of hybridization betweenO. holosericea agg. andO. scolopax agg. are discussed.