Ribosomal RNA sequence comparisons demonstrate an evolutionary relationship betweenZygnematales and charophytes

The nuclear-encoded small subunit ribosomal RNA (rRNA) sequences were determined forGenicularia spirotaenia, Mesotaenium caldariorum, andStaurastrum spec. (Zygnematales) to elucidate the evolutionary position of these green algae. Results of neighbour-joining and maximum parsimony phylogenetic analyses support a monophyletic origin of theZygnematales within the evolutionary assemblage defined by theCharophyceae (sensuMattox & Stewart) and land plants. TheZygnematales/Charophyceae/land plants are evolutionarily distinct from the monophyletic lineage defined by theChlorodendrales, Pseudoscourfieldiales, and theMicrothamniales/Chlorophyceae. In memoriamRobert W. Hoshaw.     

Relationships in theAcanthaceae and related families as suggested by cladistic analysis ofrbcL nucleotide sequences

A parsimony analysis of DNA sequences of the chloroplast-encoded generbcL from twelve members of theAcanthaceae s.l., including members of the sometimes segregateThunbergioideae andNelsonioideae, and other families in theBignoniales sensuThorne (1992) is presented. The results largely agree with the classification of theAcanthaceae presented byBremekamp (1965) andThorne (1992) and supportNelsonioideae as a sister group to the rest of theAcanthaceae. Thunbergioideae are placed as a sister toAcanthaceae s.str.Acanthus andAphelandra, both representatives ofAcanthoideae, form a sister group toRuellioideae. An analysis of branch support found that many branches throughout theBignoniales are weakly upheld. This points to the need for further studies in the group using more sequences ofrbcL as well as other data. None of the families ofBignoniales as presently circumscribed (includingAcanthaceae s.l.) were strongly supported, although the larger clade containing the families of theBignoniales was robust.     

Phylogenetic analysis ofUlmaceae

A phylogenetic analysis of theUlmaceae, Cannabaceae, Barbeyaceae, andBroussonetia of theMoraceae produced nine equally parsimonious trees with 127 steps. TheUlmoideae (Ulmaceae, sensuGrudzinskaya) are a monophyletic group and distinct from theCeltidoideae. The genusAmpelocera occupies an isolated taxonomic position among the celtidoids. The similarity ofAmpelocera to the fossil celtidoid flowerEoceltis of North America suggests thatAmpelocera posesses an archaic suite of characters, and occupies a primitive position among the celtidoids, theCannabaceae and theMoraceae. The relationships among the other celtidoid taxa,Cannabaceae, andBroussonetia are problematic. TheCannabaceae andBroussonetia of theMoraceae are nested within the celtidoids suggesting that this is a paraphyletic group. The close, but unresolved, relationship of the celtidoids to theMoraceae andCannabaceae observed in this analysis, and the appearance of the celtidoids in the fossil record prior to the ulmoids suggests that the evolutionary relationship of the ulmoids and celtidoids may be more distant than current taxonomic treatments reflect.     

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.     

Die Stipeln derIrvingioideae undRecchioideae und ihre systematische Wertung nebst Bemerkungen zur Holzanatomie und Palynologie

TheSimaroubaceae generally have no true stipules. The stipule-like appendages of some genera proved to be pseudo- or metastipules (Weberling & Leenhouts 1965). There seem to be some exceptions, however: the generaCadellia (incl.Guilfoylia) andRecchia on the one hand, and theIrvingioideae on the other. As these taxa, with exception ofRecchia, have simple leaves, there are no indications that their stipule-like appendages might be pseudo- or metastipules. In regard to their position and ontogeny these appendages behave completely like true stipules. Assuming the view ofForman, one could conceive a morphological line from the long, broadly inserted axillary stipules of mostIrvingioideae to the small scaly triangular stipules ofIxonanthoideae. The similarities between the stipules ofIrvingioideae andErythroxylaceae (already emphasized byHallier and others), become even more evident when their ontogeny is investigated. TheIrvingioideae, therefore, might be regarded as a separate family (perhaps with some relation to theErythroxylaceae,Hallier) or as a subfamily ofIxonanthaceae (Forman).—In addition to data on stipules some results on the palynology and shoot anatomy of the generaCadellia (incl.Guilfoylia) andRecchia are reported. Their relationship with theSimaroubaceae also appears doubtful. If they are to be included, they represent a somewhat isolated group near the base of the family which otherwise has lost its stipules.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Morphological phylogenetics of theAsteraceae-Asteroideae,with notes on character evolution

The relationships within theAsteraceae-Asteroideae are scrutinized by means of cladistic analyses of morphological and chemical data. New data are presented, and novel interpretations of features previously used in cladistic analyses are made. Examples are insertion of anther filaments in the corolla, and morphology of disc floret styles, respectively. The results include a polyphyletic or paraphyleticInuleae s. l., and a paraphyleticHeliantheae s.l. in which theEupatorieae constantly form a subclade. Comparisons are made with previous cladistic analyses of morphological data, and those acquired from molecular data. Speculations about the evolution of certain organs are presented.     

Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

Isoenzyme variation in the generaPhaseolus andVigna (Fabaceae) in relation to their systematics: Aspartate aminotransferase and superoxide dismutase

Evolutionary variation of aspartate aminotransferase and superoxide dismutase isoenzymes in 14 wild and cultivated species ofPhaseolus andVigna has been studied by electrophoresis and isoelectric focusing in polyacrylamide gel. The American cultivated beans of the genusPhaseolus s. str.,P. vulgaris, P. coccineus, P. lunatus andP. acutifolius, form a homogeneous group with only minor isoenzyme variation. The genusVigna, on the contrary, proves to be heterogeneous in isozyme characters. Several clusters of taxa can be distinguished in close correspondence with modern treatments of the genus. The isoenzyme data support the inclusion of the Asian Azuki beans of subg.Ceratotropis inVigna, but argue against the transfer of the S. American speciesP. adenantha. The cowpea complexV. unguiculata s. lato of sect.Catiang forms an uniform and isolated group, distinct from other sections of subg.Vigna, and shows affinity toPhaseolus s. str. by some isoenzymes. It is suggested to removeV. unguiculata s. lato from subg.Vigna and to recognize it as a separate subg.Catiang (DC.)Jaaska & Jaaska, stat. nov.     

Therps16 intron and the phylogeny of theRubioideae (Rubiaceae)

The phylogeny of the subfamilyRubioideae (Rubiaceae) was estimated from sequence variation in therps16 intron (cpDNA) in 143 ingroup and 5 outgroup taxa. The analysis largely confirms a recent one based onrbcL sequences, but branch support is often much stronger. Three of the traditional subfamilies are supported,Rubioideae, Cinchonoideae s. str., andIxoroideae s. l. while there is no support forAntirheoideae. TheRubioideae are the sister group of all otherRubiaceae and comprise the tribesAnthospermeae, Coccocypseleae, Cruckshanksieae, Coussareeae, Gaertnereae, Hedyotideae, Knoxieae, Morindeae, Ophiorrhizeae, Paederieae, Pauridiantheae, Perameae, Psychotrieae, Rubieae, Spermacoceae, Theligoneae, andUrophylleae. TheHamelieae andHillieae belong to theCinchonoideae. Rachicallis andSiemensia should be transferred from theHedyotideae to theCinchonoideae. ThePauridiantheae, Urophylleae, Ophiorrhizeae, andRaritebe form the basalmost subclade of theRubioideae. The second basalmost clade consists of the generaLasianthus andPerama. The third basalmost clade consists of the tribesCoussareeae, Coccocypseleae andCruckshanksieae, and the generaDeclieuxia andHindsia. The tribesKnoxieae, Anthospermeae, Argostemmateae, Paederieae, Theligoneae, Rubieae, Hedyotideae, andSpermacoceae are members of one clade. TheKnoxieae are monophyletic ifOtiophora, Otomeria, andPentas are included. The tribeAnthospermeae is supported as monophyletic, but its subtribes are not. ThePaederieae, together withTheligonum, form a paraphyletic grade basal to theRubieae. TheHedyotideae, includingSchismatoclada, form a grade at the base of theSpermacoceae. TheGaertnereae are monophyletic and distinct from thePsychotrieae. TheMorindeae are monophyletic and includeDamnacanthus andMitchella. Schradera is the sister group of theMorindeae. ThePsychotrieae are monophyletic when theGaertnereae, Lasianthus, andDeclieuxia are excluded. The recognition of a subtribeHydnophytineae leaves the rest of thePsychotrieae paraphyletic.Psychotria is paraphyletic with respect to all other genera of the tribe. Approximately 50 genera are here classified for the first time based on molecular data.     

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.     

Comparative ultrastructure of pyrenoids inTrebouxia (Microthamniales,Chlorophyta)

Pyrenoid ultrastructure has been investigated from cultures of all 26 species ofTrebouxia with the aim of establishing pyrenoids as a taxonomic character. Different arrangements and forms of thylakoid lamellae within the pyrenoid matrix allow eight pyrenoid types to be distinguished. Each type is characteristic of a group of species. Thegigantea- andimpressa-type are similar, differing only in the form of the tubules: short, branched tubules mark thegigantea-type; ± long and straight invaginations theimpressa-type. Thearboricola-type is characterized by meandering pyrenoid membranes developing from lamellae parallel with each other in young autospores. Pyrenoids of thegelatinosa-type are traversed by thin parallel-arranged tubules. Few thylakoids with a curved profile are typical of theirregularis-type. Thecorticola-type is different from all others in having a distinct starch sheath closely connected with the pyrenoid matrix and no pyrenoglobuli being associated with the pyrenoid membranes. No true pyrenoids have been found inT. magna andT. erici. Within the chloroplast, they have indistinct areas with pyrenoglobuli, but without differentiated thylakoids. Pyrenoid morphology is stable in culture on different media as well as in phycobionts within lichen thalli. Comparing the pyrenoid of a lichenizedTrebouxia with that from cultured species, the identification of the phycobiont within the lichen thallus is possible, without the need of culturing the algae. This has been shown in species ofParmelia andHypogymnia. New aspects for the taxonomy and systematics ofTrebouxia are discussed.Dedicated to Prof. DrLothar Geitler on the occasion of the 90th anniversary of his birthday.     

The flavonoids of theFagonia bruguieri complex (Zygophyllaceae)

Seven flavonol glycosides were identified from the main taxa of theF. bruguieri complex. Of these, kaempferol 3-rhamno-galactoside, Quercetin 3-rhamnogalactoside and Quercetin 3-galactoside are new records for the genusFagonia L. and theZygophyllaceae s. str. The distribution of these flavonoid glycosides is discussed with respect to the morphology, chemosystematics, and possible phylogeny of the complex and the genus.Dedicated to Hofrat Univ.-Prof. DrKarl Heinz Rechinger on the occasion of his 80th birthday.     

The floral nectaries in theIrvingiaceae

TheIrvingiaceae generally possess large intrastaminal receptacular disc nectaries of the mesenchymatic histo-type, which receive numerous small phloematic bundles directly from the central stele. The non-glandular epidermis bears some 10 to 15 strictly localized stomata that are deeply sunken in the parenchyma. The nectar is assumed to be exposed on the disc surface. Flowers are of simple construction, lacking specialized organs to attract pollinators. A wide range of pollinators is thus expected. TheIrvingiaceae have more characters in common withSimaroubaceae thanIxonanthaceae and should therefore be retransferred as a family of their own next toSimaroubaceae. The floral nectaries of theGeraniales and their systematic implications V. For the fourth part seeLink (1992)     

Floral structure and relationships ofHortonia (Monimiaceae)

The flowers ofHortonia angustifolia were investigated for their phyllotaxis, morphology, anatomy and development of the perianth, androecium and gynoecium. Certain features were also studied inH. ovalifolia. Characters so far overlooked further support the isolated and intermediate position of the genus between theAtherospermataceae andMonimiaceae s. str. and its archaic position among theLaurales. Dedicated to Professor Dr.W. Leinfellner on the occasion of his 70th birthday.     

Systematic relationships of theSaxifragales revealed by serological characteristics of seed proteins

Comparative investigations of serological characters of seed proteins from taxa regarded as members of theSaxifragales result in the recognition of two distinct groups of related families. One consists of theSaxifragaceae, Grossulariaceae, andCrassulaceae; to itHamamelis, and possiblyPenthorum and theRosaceae may be connected. The second group contains theHydrangeaceae, Escalloniaceae, Roridulaceae, Cornaceae, andCaprifoliaceae; it is rich in iridoid compounds, has more derived morphological characters, and seems to represent a monophyletic line block. ThePittosporaceae were not found to be linked with either of the two groups, but rather show similarities with members of theApiales. All these data support systematic arrangements proposed byDahlgren (1975a).     

Biostatistical studies on western EuropeanDactylorhiza (Orchidaceae)—theD. maculata group

Multivariate analysis tools are exploited on a data set composed of quantitative characteristics collected on 35 populations of plants of theDactylorhiza maculata (L.)Soó group from Western-Europe. These samples lead to four well-defined clusters; this, together with qualitative, cytological and ecological arguments, allows for the recognition of four specific entities:D. maculata s.str.,D. fuchsii (Druce)Soó,D. saccifera (Brongn.)Soó andD. caramulensis (Vermeulen)Tyteca. It is concluded that the floral characters play an essential role in the taxonomical distinction. It also appears that the set of characters measured, as well as the methods exploited, are especially well-suited and valuable tools for the morphological study of the genusDactylorhiza.     

Vergleichende cytotaxonomische Untersuchungen anRanunculus seguieri und der Artengruppe desR. alpestris (Ranunculaceae)

Morphological and cytological investigations as well as crossing experiments were carried out withRanunculus seguieri Vill. and 4 species of theRanunculus alpestris L. group (R. alpestris L.,R. traunfellneri Hoppe,R. bilobus Bertol.,R. crenatus Waldst. & Kit.). ForR. seguieri andR. alpestris, localities and distribution are given in addition to extensive diagnoses. A key to the species includes morphological characteristics and distribution data forR. traunfellneri, R. bilobus, andR. crenatus. New diagnostic characters are described. Crossing experiments betweenR. seguieri and the species of theR. alpestris group were unsuccessful. All 5 species have a chromosome number of 2n = 16, the record forR. bilobus is new. There is no statistically significant difference between the karyotypes ofR. seguieri andR. alpestris s. str. Nevertheless, according to morphological evidence and crossing experiments,R. seguieri is not closely related to theR. alpestris group.

     

Sind Farne Kormophyten? Eine Alternative zur Telomtheorie

Modern ideas concerning cormophyte phylogeny are strongly influenced by the telome theory. Erect growth, radial symmetry, dichotomy, protostely and eusporangial structure are considered to be primitive features. Yet, the cormus needs redefinition. TheHypolepidaceae (s. str.) are shown to have rather a two-dimensional thallus than a three-dimensional cormus. Their so-called rhizome develops a continuous areophore margin which is connected with the margin of the fronds. The longitudinally inserted fronds and branches are produced by the terminal rhizome meristem which itself grows in many respects like the apical frond meristem. The rhizomes ofDennstaedtiaceae and theDavallia-type of rhizomes are intermediate between thisHypolepis-type and the true cormus-type. TheHypolepis-type is compared with fern prothalli. Under perhumid tropical conditions higher land plants may have evolved from small, creeping, two-dimensional fern prothallium-like progenitors with isomorphous gameto- and sporophytes.

Vorgetragen auf der Morphologentagung in Zürich 1975.     

Omphalotaceae fam. nov. undPaxillaceae,ein chemotaxonomischer Vergleich zweier Pilzfamilien derBoletales

TheOmphalotaceae fam. nov., which include the generaOmphalotus andLampteromyces, are defined on the basis of characteristic sesquiterpenes and of their ability to cause white-rot. Anatomical and morphological features of the representatives of these genera support the creation of this new family. The occurrence of pigments, typical of theBoletales, and of cyanophilous spores, indicate membership of theOmphalotaceae in the orderBoletales. Relationships to the other families of this order — especially to thePaxillaceae — are discussed. The possible functional significance of fungal metabolites is considered.

Herrn Prof. Dr.J. Poelt zur Vollendung seines 60. Lebensjahres gewidmet.—Veränderte Fassung eines Vortrages auf der Tagung der Deutschen Botanischen Gesellschaft, Wien, September 1984.     

Interspecific hybridizations among species of theElymus semicostatus andElymus tibeticus groups (Poaceae)

Interspecific hybridizations were made between species of theE. semicostatus group, viz.,E. semicostatus (Nees exSteud.)Meld.,E. validus (Meld.)B. Salomon,E. abolinii (Drob.)Tzvel., andE. fedtschenkoi Tzvel., and species of theE. tibeticus group, viz.,E. pendulinus (Nevski)Tzvel.,E. tibeticus (Meld.)Singh,E. shandongensis B. Salomon, andE. gmelinii (Ledeb.)Tzvel., as well as among species within theE. tibeticus group. All species are tetraploid (2n = 4x = 28) and possess SY genomes. Meiotic pairing data from 24 hybrids involving 17 interspecific combinations are presented. The average number of chiasmata per cell ranged from 17.91 to 26.20 in hybrids within theE. tibeticus group, compared with 7.26 to 22.04 in hybrids between the two species groups. Despite the extensive collection of cytological data, there was no definite evidence for confirming or disproving the separate existence of the two groups.