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.     

Ultrastructure of pollen exine in centrospermous families

Transmission electron microscopy was utilized to examine pollen walls of selected taxa in theAizoaceae, Amaranthaceae, Basellaceae, Cactaceae, Chenopodiaceae, Didiereaceae, Halophytaceae, Nyctaginaceae, Phytolaccaceae, Portulacaceae, andCaryophyllaceae. This conspectus is an adjunct to scanning electron microscope observations of pollen surfaces and is directed towards elucidating the basic wall structure for these families. Although differences in internal morphology were observed at the inter- and intra-familial levels, they were interpreted as reflecting variations rather than major differences. The data indicate close morphological similarities of the first ten families enumerated above, i.e., those containing betalains. TheCaryophyllaceae, an anthocyanin family, indicated a slightly greater heterogeneity of pollen ultrastructure but not to the extent of disassociating it from the betalain families. In fact, this heterogeneity was rivaled by comparable heterogeneity among and within some of the betalain families. The conclusion is that all families have close pollen morphological relationships.Presented in the Symposium Evolution of Centrospermous Families, during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

Serologische Untersuchungen zur Gliederung derBrassicaceae

The serological investigations support the opinion ofJanchen (1942) to combine the generaBunias, Isatis, andSisymbrium in the tribeSisymbrieae; Cheiranthus, Erysimum, andMatthiola in the tribeHesperideae; andBrassica, Crambe, Sinapis, andSuccowia in the tribeBrassiceae. They further underline the central position of theSisymbrieae and the isolated position of theHeliophileae. In accordance withEigner (1973) theBrassiceae are placed closer to theSisymbrieae than inJanchen; the same holds for thePringleeae. No serological justification could be found to uniteArabis andBarbarea in the tribeArabideae, andAlyssum andLunaria in theAlysseae. From the antigen-systems used among the representatives ofJanchen's Lepidieae the generaLepidium andNeslia show remarkable correspondence both toCamelina andThlaspi, but not toCochlearia which appears distant fromCamelina andThlaspi also.

Teil 1/Part 1.     

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.     

Pigment dichotomy and DNA-RNA hybridization data for centrospermous families

The structural types, biogenesis and distribution of betalains and anthocyanins in centrospermous families are reviewed. The implications of the pigment and DNA-RNA hybridization data, along with other evidence, are discussed with respect to the view that the evolutionary line of centrospermous families contains eleven basic or core families of which nine produce betalains and two, theCaryophyllaceae andMolluginaceae, produce anthocyanins; it is suggested that the betalain and anthocyanin families developed from a common ancestor at a time prior to the widespread occurrence of floral pigments in angiosperms.Presented in the Symposium Evolution of Centrospermous Families, during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

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.     

Anamylopsoraceae — a new family of lichenized ascomycetes with stipitate apothecia (Lecanorales: Agyriineae)

The anatomy, chemistry and developmental morphology ofAnamylopsora pulcherrima is investigated. Some characters, including the ascus structure, suggest a close affinity with theAgyriaceae. However, the chemistry and the pycnidial structure differ as well as the ascoma ontogeny.Anamylopsora has a gymnocarpous ascoma development and the ascogonia are produced in stipes.Trapelia coarctata, as a typical member of theAgyriaceae, shows a hemiangiocarpous ascoma ontogeny. The anatomical, chemical and ontogenetical characters of several families are compared withAnamylopsora and it is shown that the genus is best placed in a monotypic familyAnamylopsoraceae Lumbsch & Lunke, fam. nova, which is placed in theAgyriineae (Lecanorales).This paper is dedicated to Prof. DrAino Henssen (Marburg) on the occasion of the 70th birthday.     

The application and citation of the generic nameParmeliopsis (Lecanorales,Parmeliaceae)

The nomenclatural history of the generic nameParmeliopsis is reviewed. Its correct citation is found to beParmeliopsis (Nyl. exStizenb.)Nyl., dating from 1866, not 1869 as commonly cited, withP. placorodia (Ach.)Nyl. as holotype species. There is consequently no nomenclatural problem to the adoption ofForaminella Fricke Meyer, typified byF. ambigua (Wulfen)Fricke Meyer for theParmeliopsis species with falcate conidia. A synopsis of the nomenclature of the North American and European species of both these genera is included.     

The phylogenetic relationships ofByblis andRoridula (Byblidaceae-Roridulaceae) inferred from partial 18S ribosomal RNA sequences

The phylogenetic relationships of the angiosperm generaByblis andRoridula have been the subject of ongoing taxonomic controversy. Twenty-eight taxa of varying degrees of alleged relationship, including 3 members of theWinteraceae (as an outgroup), were investigated using partial sequences of 18S rRNA (small subunit) and also compared against the morphological data set fromHufford's (1992) cladistic treatment of 80 members of theRosidae-Asteridae. The morphological analysis placed the two genera in a clade with theSarraceniaceae in theCorniflorae-Asterid group as a sister taxon to anEricales-Hydrangeales clade. The 18S rRNA analysis supports the recently publishedrbcL DNA analysis ofAlbert & al. (1992), withRoridula joined to taxa in the lowerRosidae, butByblis joining instead to members of theAsteridae near theSolanaceae. Comparisons for congruence between the three analyses placeByblis in the higher Asterid group near theSolanaceae, andRoridula possibly nearer theSarraceniaceae andEricales. These results imply that the traditional morphological characters used to relate the two genera are possibly the result of convergence towards similar ecological and life-history strategies rather than synapomorphies.     

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).     

Phylogenetic relationships of theJuglandaceae

A cladistic analysis of molecular data from the chloroplast generbcL was used to examine the taxonomic relationships of the walnut family (Juglandaceae). In addition, chemical and morphological data from a previous study byHufford (1992) were incorporated, expanded, and analyzed independently and in combination with the molecular data. The results of these analyses suggest that theJuglandaceae are more closely related to theFagaceae, Betulaceae, Casuarinaceae, andUrticaceae and their relatives (sensuCronquist 1981) than they are to theAnacardiaceae (sensuThorne 1983). However, sequence data fromrbcL also suggest a relationship between the higherHamamelidae and certain families in theRosidae sensuCronquist 1981 (such asRosaceae andRhamnaceae), an outcome which would add credence to the widely accepted view of the polyphyletic nature of theHamamelidae.     

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.     

Contributions to the knowledge of sieve-element plastids inGunneraceae and allied families

P-type sieve-element plastids were found in theGunneraceae, while S-type plastids are present in theHaloragaceae andHippuridaceae. The specific characters of the sieve-element plastids (e.g., their size and the morphology of their contents) are discussed in relation to other taxa of theRosidae containing P-type plastids and to the systematic position of theGunneraceae. Contributions to the Knowledge of P-Type Sieve-Element Plastids in Dicotyledons, III. — For other parts of this series see (I.:)Behnke (1982 b) and (II.:)Behnke (1985).     

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.

     

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.     

Chemosystematics ofSuillaceae andGomphidiaceae (suborderSuillineae)

The chemotaxonomic findings relating to the generaBoletinus, Suillus, Gastroboletus, Gomphidius, andChroogomphus are summarized and discussed, using published data as well as our own hitherto unpublished evidence of pigments and chromogens. The study confirms repeatedly made claims that these genera are closely related. In addition to the presence of pigments which are typical for most members of theBoletales (e.g., pulvinic acid derivatives, terphenyl quinones, cyclopentenones), prenylated phenols and quinones can also be constantly detected here (with the exception ofBoletinus), just as inRhizopogon. Accordingly,Suillus is more closely related to theGomphidiaceae andRhizopogonaceae than to the remaining boletes. It is therefore necessary to establish a new family (Suillaceae which includeBoletinus, Suillus, andGastrosuillus) and a new suborder (Suillineae which includeSuillaceae, Gomphidiaceae, andRhizopogonaceae) within theBoletales. Chemosystematics ofBoletales 2. For part 1 seeBesl & al. (1986). Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Chromosome numbers and differentiation of centrospermous families

The patterns of variation of chromosome numbers in theCentrospermae suggest a common ancestry of centrospermous anthocyanin and betalain families. Phylogenetic divergence in the order may have originated with progenitors similar to extantMolluginaceae, Aizoaceae orPhytolaccaceae taxa with x = 9. Evolutionary radiation and advancement in several lines then seems to have been paralleled by trends for increasing chromosome base numbers through dysploidy and polyploidy, e.g. towardsCaryophyllaceae, Portulacaceae-Basellaceae, Hectorellaceae, Cactaceae, Didieraceae, Nyctaginaceae andChenopodiaceae-Amaranthaceae. Presented in the Symposium Evolution of Centrospermous Families, during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

Studies on the early floral development inCleomoideae (Capparaceae) with emphasis on the androecial development

Floral ontogeny ofCleome spinosa, Cleome violacea andPolanisia dodecandra subsp.trachysperma was studied in the context of the question whether the fascicled androecium ofReseda andCapparis (with fused fascicles) or the 2 + 4-pattern of theBrassicaceae is primitive in theCapparales. InPolanisia dodecandra, the 9–18 stamens show unidirectional initiation from the adaxial side toward the abaxial side of the flower. InCleome violacea, the six stamens also are formed in an unidirectional order, but development starts abaxially and a zigzag-like pattern is superimposed. InCleome spinosa, two stamen primordia in transversal (lateral) position are followed by four stamens which arise on a somewhat higher level in two pairs in front of the median sepals. It is assumed that the evolutionary steps in the androecial development proceed fromReseda viaCapparis andPolanisia/Cleome toBrassicaceae. This interpretation is supported byrbcL-studies (Chase & al. 1993,Rodman & al. 1993).Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

P-type sieve-element plastids present in members of the tribesTriplareae andCoccolobeae (Polygonaceae) renew the links between thePolygonales and theCaryophyllales

Form-Pfs sieve-element plastids were found inTriplaris, Ruprechtia, andCoccoloba (Polygonaceae) while other genera of the family and those studied from the often associatedPlumbaginaceae contain S-type sieve-element plastids. The rareness of form-Pfs plastids among the angiosperms, their similarity to the peculiar form-P3fs plastids of theChenopodiineae, and the comparatively small plastid diameters measured for all forms present in theCaryophyllales, Polygonales, andPlumbaginales suggest close relationships between these taxa. The restriction inPolygonaceae of form-Pfs plastids to the closely allied tribesTriplareae andCoccolobeae is discussed with regard to both the intrafamilial and ordinal phylogeny, and also considering possible connections to the only magnoliidaean Pfs-taxonCanella. Dedicated to Univ.-Prof. DrF. Ehrendorfer on the occasion of his 70th birthday.     

The genusAntennaria (Asteraceae: Inuleae) in western North America: Morphometric analysis ofAntennaria alborosea,A. corymbosa,A. marginata,A. microphylla,A. parvifolia,A. rosea,andA. umbrinella

Multivariate analysis of vegetative and reproductive characters was used to examine morphological relatedness amongAntennaria alborosea A. E. Porsild,A. corymbosa E. Nels.,A. marginata Greene,A. microphylla Rydb.,A. parvifolia Nutt.,A. rosea Greene, andA. umbrinella Rydb. Both pistillate and staminate plants were examined. Some of the characters examined were variable in one species, but stable in another (i.e., presence or absence of papillae on the achenes). Our analyses indicate that the seven species are morphologically distinct. It is hypothesized that theA. rosea agamic complex arose through hybridization amongA. corymbosa, A. microphylla, A. umbrinella, and possiblyA. dioica (L.)Gaertn. However, hybridization between the three former species and others, as well as their subsequent morphological responses to different environmental conditions causes confusion in recognizing the taxa.Antennaria angustifolia Rydb.,A. arida E. Nels.,A. confinis Greene,A. scariosa E. Nels.,A. foliacea humilis Rydb.,A. concinna E. Nels., andA. viscidula E. Nels. are considered to represent F 1 hybrids.