Delimitation ofPelargonium sect.Glaucophyllum (Geraniaceae)

Pelargonium otaviense Knuth andP. spinosum Willd. are excluded from sect.Glaucophyllum, whileP. grandiflorum (Andr.)Willd.,P. patulum Jacq. andP. tabulare (Burm. f.)L'Hérit. of sect.Eumorpha are included. Sect.Glaucophyllum is characterized by green to glaucous vegetative organs and zygomorphic white to pink corolla with five narrow petals. All the species have an identical pollen and chromosome morphology, the same basic chromosome number (x = 11) and similar flavonoid patterns. A close relationship between sect.Glaucophyllum and sect.Pelargonium is indicated by the occurrence of natural hybrids and concordant characters. Isorhamnetin and luteolin have been detected in the genus for the first time.     

A cytological study ofPelargonium sect.Eumorpha (Geraniaceae)

The chromosome numbers of seven species ofPelargonium sect.Eumorpha have been determined from material of known wild origin, and karyotypic comparisons have been made. Within the section there is variation in basic chromosome number (x = 4, 8, 9, 11), variation in chromosome size, and two species have polyploid races. The three species with chromosome numbers based on x = 11 have the smallest chromosomes (1.0–1.5 µm); chromosomes are larger (1.0–3.0 µm) in the other species.P. elongatum has the lowest chromosome number in the genus (2n = 8).P. alchemilloides is exceptional in that it has four cytotypes, 2n = 16, 18, 34 and 36, and the form with 2n = 36 has large chromosomes (2.0–5.0 µm). Evidence from a synthesized hybrid suggests thatP. alchemilloides with 2n = 16 may be of polyploid origin. The three species based on x = 11 appear to be more closely related to species from other sections ofPelargonium that have the same basic chromosome number and small chromosome size, rather than to other species of sect.Eumorpha.     

A reappraisal ofPelargonium sect.Ligularia (Geraniaceae)

A study of 33 species ofPelargonium sect.Ligularia reveals four basic chromosome numbers, x = 8, 9, 10, and 11, and variation in chromosome size. From evidence of karyology and hybridization attempts, proposals are made to divide the section into smaller groups and to transfer some species to other sections.     

A trnL-F based phylogeny for species ofPelargonium (Geraniaceae) with small chromosomes

Phylogenetic analysis was performed of 921 positions of trnL (UAA) 5 exon — trnF (GAA) exon chloroplast DNA regions from 68 representatives ofPelargonium sectt.Campylia, Cortusina, Glaucophyllum, Hoarea, Isopetalum, Ligularia, Otidia, Pelargonium, Peristera, Polyactium, andReniformia, together with five putative outgroup species from sectionsCiconium, Chorisma andJenkinsonia. The total data set therefore comprised 67.2 kb of DNA sequence. Two main ingroup clades were identified: one clade contains sectionsPeristera, Reniformia, andIsopetalum, the other contains sectionsCampylia, Cortusina, Glaucophyllum, Hoarea, Ligularia, Otidia, Pelargonium, Polyactium and two species currently grouped in sect.Peristera. Branching order among five main clades within the latter clade was not resolved. The trnL-F sequence data support monophyly only for sectionsReniformia andHoarea, the remainder of the currently recognized sections ofPelargonium being either paraphyletic or polyphyletic. The data further suggest that sect.Polyactium is diphyletic and that sect.Glaucophyllum is nested within sect.Pelargonium. One relatively derived clade, which represents half of the genus, contains predominantly geophytic and succulent species, occurring in the geographically restricted winter rainfall region of the South African Cape. This pattern is interpreted as reflecting explosive radiation, possibly as an adaptive response to recent aridification in the western Cape.     

Untersuchungen zur Karyologie und Mikrosporogenese vonPelargonium sect.Pelargonium (Geraniaceae)

The chromosome numbers of the 24 species of sect.Pelargonium were determined from field collected and cultivated plants of known localities in S. Africa. Twelve species are diploid (2n = 22), eight tetraploid (2n = 44), one hexaploid (2n = 66), and three octoploid (2n = 88). The chromosome numbers correlate well with the proposed subdivision of sect.Pelargonium. Its chromosomes are relatively small (1.0–1.5 µm) in comparison to most of the other sections, and its diploid karyotype is considered to be primitive. The occurrence of the basic number x = 11 in this section, in other sections of the genus, and in related genera (Monsonia, Sarcocaulon) leads to the conclusion that x = 11 probably is basic for the whole genus. — The pollen meiosis, microsporogenesis and pollen fertility of the diploid species is normal, with the exception of one, possibly young taxon from the Greyton Nature Reserve. The tetraploid species could be of autoploid origin, the higher polyploids exhibit a mixed auto-alloploid nature. — The 20 diploid and tetraploid species have a relatively small distribution range, most of them occur in the SW. Cape Province of South Africa. This area may therefore be considered as the centre of origin of the genus. Three of the four high polyploid species occupy rather large areas.

Untersuchungen zur Karyologie und Mikrosporogenese der GattungPelargonium, 1.     

Phylogenetic relationships withinPelargonium sect.Peristera (Geraniaceae) inferred from nrDNA and cpDNA sequence comparisons

Phylogenetic analysis of nrDNA ITS and trnL (UAA) 5 exon-trnF (GAA) chloroplast DNA sequences from 17 species ofPelargonium sect.Peristera, together with nine putative outgroups, suggests paraphyly for the section and a close relationship between the highly disjunct South African and Australian species of sect.Peristera. Representatives fromPelargonium sectt.Reniformia, Ligularia s. l. andIsopetalum (the St. Helena endemicP. cotyledonis) appear to be nested within thePeristera clade. The close relationship between the South African and AustralianPeristera is interpreted as being caused by long-range dispersal to Australia, probably as recent as the late Pliocene.     

Additional chromosome numbers inPelargonium (Geraniaceae)

Mitotic chromosome counts were made from field collected and subsequently cultivated plants of 61Pelargonium species from 14 sections. The 33 new results are presented. 47 of the species have a basic number of x = 11, nine spp. of x = 9 and five spp. of x = 8. 17 spp. are polyploid. In two sections species with different basic numbers occur, which is of interest for the subgeneric classification. The size of the chromosomes varies between the investigated species. Most but not all species with x = 11 have short, those with x = 8, 9 large, and only exceptionally short chromosomes. Within sections chromosome size is not always uniform. The relationship between the different basic chromosome numbers is discussed.     

Karyological studies inPelargonium sectt.Ciconium,Dibrachya, andJenkinsonia (Geraniaceae)

Pelargonium sect.Ciconium and sect.Dibrachya have a basic chromosome number of x = 9, whereas sect.Jenkinsonia has x = 9, 11, and 17.     

Taxonomic revision of Geranium sect. Gracilia (Geraniaceae)

Geranium sect.Gracilia (Geraniaceae) comprises nine species in the Andes of Venezuela and Colombia.Geranium lindenianum andG. gracilipes are synonymized withG. holosericeum. The identity ofG. multiceps is clarified. Its differences from the two Colombian species,G. santanderiense andG. lainzii, are discussed.Geranium lignosum, G. sebosum, G. stoloniferum, G. subnudicaule, andG., velutinum are now considered to be members of sect.Gracilia. The identity ofG. pilgerianum could not be clarified because the type was destroyed and other collections have not been found. Micromorphological leaf and seed characters are discussed. Six lectotypes and one neotype are designated.

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Resumen  Se revisaGeranium sect.Gracilia (Geraniaceae), un grupo formado por nueve especies distribuidas por los Andes de Venezuela y Colombia. Se sinonimizanG. lindenianum y G. gracilipes aG. holosericeum. Se aclara la identidad deG. multiceps y se establecen sus diferencias conG. santanderiense y conG. lainzii, ambas esencialmente colombianas. Se aceptan en esta secciónG. lignosum, G. stoloniferum yG. sebosum, propuestos tras la descripción de la sect.Gracilia, yG. subnudicaule yG. velutinum, ya tratados por otros autores en como pertenecientes a este grupo. Por el contrario, la identidad deG. pilgerianum no ha podido ser aclarada tras la destrucción del tipo y ante la falta de otros materiales de la especie. Se estudian diversos caractere micromorfológicos, especialmente algunos referibles a la anatomía foliar y la de la semilla. Se designan seis lectótipos y un neótipo.

     

The chemotaxonomy ofGeranium (Geraniaceae)

Relationships amongGeranium species constituting sectt.Anemonifolia, Lucida (monotypic) andRuberta together with representatives of sect.Unguiculata were investigated by gas-chromatographic study of essential oils, electrophoretic comparison of seed proteins and chromatographic separation of nectar amino acids. — Essential oil study gave little information.G. macrorrhizum (sect.Unguiculata) had far greater quantities of essential oils in its foliage than other species and differed from them qualitatively. — Species of sectt.Anemonifolia andRuberta, together withG. cataractarum (sect.Unguiculata), between them yielded 19 seed protein bands; the distribution of these indicated close relationship among the species and was consistent with hypotheses for the origin of certain species by alloand autopolyploidy partly within the group. Involvement of an unknown species in the origin of two allopolyploids was implied. Separate origins for the two octoploid species in this set are also inferred. In two instances there was evidence for the transformation of one band into another subsequent to the separation of related species. The inference of allopolyploidy was supported by the occurrence of additivity of parental bands shown by an artificial hybrid between two of the species. A model for the evolutionary divergence of the seed protein patterns is presented. Two species outside the above set appeared less closely related; they wereG. lucidum andG. macrorrhizum (sect.Unguiculata) and between them they showed 6 additional bands, four of which were shared. — Of 18 nectar amino acids found, 4 to 15 occurred in any one species, with low numbers (4 and 8) occurring in the most extreme inbreeders. The spectra of nectar amino acids of two hybrids showed additivity of those of the respective parents. The results echo rather closely those provided by seed proteins, but in the absence of data from outside the group their taxonomic significance is uncertain. However, the divergence between the two octoploids was again evident.     

Wuchsform und Organisation vonMonsonia nivea (Geraniaceae) unter besonderer Berücksichtigung der Blattstieldornen

Monsonia nivea is a hemicryptophyte which shows spiny petioles after the loss of leaf blades. Anthocladial inflorescences and their supporting vegetative parts are spineless. Spines are formed only by vigorous rosettes. Similar spiny petioles can be found in all other genera of the tribeGeranieae.

     

Pelargonium caroli-henrici (Geraniaceae), a new species from the Western Cape Province

Pelargonium caroli-henrici spec. nova (sect.Hoarea) is described. The new species is apparently endemic to quartzite areas of Vanrhynsdorp Division in the western Cape Province (South Africa). The chromosome number 2n = 22 was counted on specimens in cultivation.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

Systematic revision ofErodium (Geraniaceae) in Egypt

A key is provided for the 14Erodium species of the Egyptian flora. The important differential chracters of leaf, inflorescence, flower, and fruit are discussed and illustrated.Systematic Revision ofGeraniaceae in Egypt, I.     

Cytological variation and evolution withinPelargonium sectionHoarea (Geraniaceae)

Chromosome numbers of 65 species of sect.Hoarea have been determined. These show three basic chromosome numbers, x = 11, 10 and 9. Only a few species are tetraploid. In five species both diploid and tetraploid cytotypes are reported. Several cases of deviations in chromosome numbers and cytological abnormalities were found, most of these being related to the presence of B chromosomes that occur in eight species. Evidence is presented to suggest that the basic chromosome numbers of x = 10 and x = 9 are derived from x = 11 by centric fusion. Although variation in basic chromosome number withinPelargonium has been the subject of detailed study, this is the first time that evidence has been found for a mechanism of change in basic number, that of centric fusion by Robertsonian translocation. For the species of sect.Hoarea with x = 9, where the evidence for Robertsonian translocation is greatest, this process has probably taken place quite recently. In contrast to results from other sections of the genusPelargonium, the three different basic numbers of sect.Hoarea do not contradict its delimitation as a natural taxon.     

Karyological and chemotaxonomical studies of the hybridPelargonium incarnatum ×P. patulum (Geraniaceae) and its parents

Pelargonium incarnatum is a tetraploid species (2n = 40) andP. patulum is diploid (2n = 22). A natural hybrid, collected at the same locality as the parent species, has 2n = 42 chromosomes. The meiosis of the hybrid is practically normal and its pollen fertility corresponds to that ofP. incarnatum. The flavonoid patterns of the parent species and the hybrid are very similar. The origin and phylogenetic implications of this intersectional hybrid are discussed.     

Biosystematic studies inTulipa sect.Eriostemones (Liliaceae)

Evolutionary relationships and infra- and interspecific variability within the species ofTulipa sect.Eriostemones Boissier were studied. Measurements of 35 morphological characters were used for principal component and canonical variate analyses. Crossing experiments and chromosome counts were carried out. Two axes of morphological variation were encountered within the principal component plot. Variation of subsect.Biflores was traced along one axis, the species of subsect.Australes as well asSaxatiles were found along the other axis. The nomenclature of names of subsect.Biflores was considered on the basis of their geography, specific morphological characteristics and polyploidy. Conspecificity of names of all species was tested based on both morphology and crossability.     

Revision ofVerbesina sect.Pseudomontanoa (Asteraceae)

Verbesina sect.Pseudomontanoa is revised. The last treatment of the group byRobinson & Greenman (1899) recognized 5 species; the present treatment recognizes 12 species, 3 of which (V. breedlovei, V. cronquistii andV. olsenii) are described as new. A key to species, phyletic diagram and distribution maps are provided.     

Fertility relationships ofGeranium (Geraniaceae): sectt.Ruberta,Anemonifolia, Lucida andUnguiculata

Cross-pollinations were carried out among 11 briefly described species ofGeranium. Eight species pairs produced hybrids, of which five had not been reported before. The close relationship ofG. purpureum, G. robertianum andG. rubescens (sect.Ruberta) was confirmed; they form a polyploid series (diploid, tetraploid and octoploid on base x = 16). ForG. canariense (sect.Anemonifolia), another octoploid on base x = 16, the results suggest greater affinity with the former section.G. maderense andG. palmatum of sect.Anemonifolia (2n = 68) are confirmed as closely related to each other.G. maderense produced hybrids withG. robertianum (2n = 64; sect.Ruberta) and withG. cataractarum (2n = 36; sect.Unguiculata). Meiosis in the latter hybrid suggests allopolyploidy between parents with 2n = 32 and 2n = 36. Whereas all these species clearly form a very close alliance,G. lucidum (sect.Lucida) andG. macrorrhizum andG. dalmaticum (both sect.Unguiculata), appear genetically more isolated from them. One plant ofG. macrorrhizum ×G. dalmaticum was raised. — In crosses where hybrids or non-germinating seeds resulted, the reciprocal cross in the majority of cases produced a greatly inferior result or none at all. This asymmetry of response could in some cases be explained by inability of pollentubes from short-styled parents to reach the ovary of a long-styled species and in others by a modification of Hogenboom's theory of incongruity, but neither explanation works for every case. In all our asymmetric results where the ploidy level differed the diploid was the successful female, not the tetraploid, as is usually the case. — Some variation in results from year to year could be attributed to weather conditions.Dedicated to Hofrat Univ.-Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

Floral divergence and convergence in the genusPelargonium (Geraniaceae) in southern Africa: Ecological and evolutionary considerations

Based on field observations and a survey of the available literature, the functional and evolutionary significance of floral characters ofPelargonium is investigated in relation to a recent infrageneric re-classification. Most of the 208Pelargonium taxa (recognized as species, subspecies or varieties) involved show bee and long-proboscid hovering fly pollination syndromes (about 60% and 25%, respectively), only 7% of the taxa are pollinated by butterflies, some 2 to 4% by hawkmoths and presumably 1% by birds. The heterogeneity ofPelargonium in terms of structural blossom types and pollination syndromes indicates an independent and repeated evolution of convergent flower morphs in the genus and even in sections.     

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.