Implications of morphological phylogenetics for the placement of the generaAdenocaulon andEriachaenium (Asteraceae)

Adenocaulon andEriachaenium are two problematic genera because their tribal and subfamilial placement in Asteraceae is uncertain. Previous cladistic analyses based on molecular data undertaken to analyze the relationships within Asteraceae, placeAdenocaulon in the tribe Mutisieae (Cichorioideae). This paper investigates cladistic relationships amongAdenocaulon andEriachaenium and tribes of subfamilies Cichorioideae and Asteroideae using morphological data. Thirty-eight characters were scored across 52 genera selected as exemplar taxa to represent the current classification system. In the analysis (one tree, length = 86, c.i. = 0.55, r.i. = 0.64)Adenocaulon andEriachaenium are sister taxa and appear as an isolated clade nested in Cichorioideae. A new, tentative position among the tribes of the paraphyletic Cichorioideae is proposed for these two isolated genera.     

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.     

Chromosome numbers and rust susceptibility as taxonomic criteria inRosaceae

BothNeuradoideae andChrysobalanoideae seem rust-free. TheSpiraeoideae andPomoideae are heavily susceptible toGymnosporangium rusts. While thePrunoideae resemble theRosoideae in being vulnerable to attacks ofPuccinia species, they are additionally susceptible toTranzschelia andThekopsora, whereas theRosoideae are characteristically afflicted byPhragmidium.—It is suggested (a) to treat theChrysobalanoideae as a separate family (Chrysobalanaceae), (b) to transferDryas from theRosoideae-Potentilleae-Dryadinae to theRosoideae-Cercocarpeae, and (c) to divide theRosoideae into two main groups of tribes: (i) the rust-freeKerrieae andCercocarpeae with x = 9, and (ii) the rust-susceptiblePotentilleae, Ulmarieae, Roseae andSanguisorbeae with x = 7.     

Cladistic analyses of some net-veinedLiliiflorae

Comparisons between previous intuitive or phenetic classifications of the netveinedLiliiflorae was used to determine a suitable group of taxa for cladistic analyses. The resultant 48 genera and generic groupings comprising 19 families from within theAsparagales, Liliales andDioscoreales were then analysed. The cladistics largely supported previous treatments, with theLiliales, Dioscoreales andAsparagales all monophyletic.Invited paper, 14th International Botanical Congress, Berlin, Symposium on theory and practice of botanical classification, July 1987.     

Chloroplast DNA reassociation and grass phylogeny

Sequence variation in chloroplast DNA (cpDNA) as measured by DNA reassociation was examined in 12 grass species to address systematic problems in thePoaceae at the subfamilial and tribal levels. Two species,Petunia (Solanaceae) andGlycine (Leguminosae), were included to determine degrees of sequence divergence in cpDNA between monocots and dicots. The data were analyzed phenetically and phylogenetically. Species were segregated into four major groups that corresponded to the subfamiliesPooideae, Oryzoideae, Chloridoideae, andPanicoideae. Representatives of thePooideae andOryzoideae grouped together as did members of theChloridoideae andPanicoideae. ThePooideae split into two major groups corresponding to the recently recognized supertribesTriticanae andPoanae. Internodes between subfamily branches were short which might indicate a burst of divergence in the family early in its evolution. Sequence similarity values between the monocot grass species and the two dicot taxa ranged from 0.15 to 0.27, representing the highly conserved sequences of the chloroplast genome.     

Accumulation of seed storage proteins and the taxonomy ofPoaceae

The accumulation of specific seed proteins is a taxonomically valuable feature and can be used to additionally characterize plant taxa. To date, mainly crop proteins have been analysed in thePoaceae. In this investigation seed proteins from 147 species were screened with emphasis on legumin-like proteins and prolamins. The groups resulting from evaluation of the protein profiles correspond with well-known subfamilies and tribes.Panicoideae are clearly separated fromPooideae. WithinPooideae, theBromeae plusTriticeae tribes revealed obvious similarities.Lolium, Festuca andVulpia, generally included in the tribeFestuceae, revealed a protein profile similar to the profile of theBromeae/Triticeae. Legumin-like proteins are accumulated abundantly inBambusoideae andPooideae exceptBromeae/Triticeae, however, only the species included in theAveninae subtribe produce soluble (globulin-type) legumins as already known fromAvena sativa. Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Cytoevolutionary patterns inRutaceae

Chromosome numbers for 9 tribes and 73 genera ofRutaceae are examined for the probable chromosome base numbers in these taxa. There is abundant dysploidy and infrageneric polyploidy in the largeRutoideae/Toddalioideae complex. We found that x = 18 was typical for the tribesZanthoxyleae andToddalieae; probably ancestral in theBoronieae and perhaps in theRuteae, Diosmieae, andCusparieae; and characteristic of subfamilyFlindersioideae. Considering the basic position of elements ofZanthoxyleae andToddalieae in the family it appears that diploid x = 18 is ancestral in theRutaceae. The morphologically advancedCitroideae are invariant for x = 9 and may be a product of dysploid reduction.     

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.     

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.     

Phylogenetic relationships in theCampanulales based onrbcL sequences

Phylogenetic relationships within the angiosperm orderCampanulales were investigated by comparative sequencing of the chloroplast generbcL. CompleterbcL sequences were obtained for ten species in six families within the order. These data were analyzed along with previously publishedrbcL sequences from other taxa (for a total of 117 species) within the subclassAsteridae and outgroups, producing 32 equally parsimonious trees. A subset consisting of 44 of these taxa was then chosen and more rigorous analyses performed, resulting in four equally parsimonious trees. Results indicate that two major clades roughly corresponding to traditionally circumscribedAsterales andCampanulales exist as sister taxa. In particular, therbcL trees indicate thatSphenoclea is not a member ofCampanulales orAsterales, thatPentaphragma is more closely allied toAsterales thanCampanulales, that theCyphiaceae are not monophyletic, thatCampanulaceae andLobeliaceae are not sister taxa, and thatStylidiaceae are correctly placed withinCampanulales.     

Stomatal types ofCactaceae

The stomatal types (i.e. the arrangement of epidermal cells in the vicinity of a stomatal pore in superficial view) have been examined in more than 150 taxa ofCactaceae, mostly using documented material. Preparations have been made by mazerating pieces of tissue with modified Jeffrey's Solution and staining with chlore-zinc-jodine.—The examined members of the subfamilyCactoideae showed parallelocytic stomata with only minor deviations in a number of cases. Members of the subfamiliesPereskioideae andOpuntioideae show parallelocytic stomata on the leaves, but the stomatas of the stem represent a different type, which apparently is not yet described. It is termed opuntioid here. In most cases the stomata are superficial but a few taxa show markedly sunken or hidden stomata.—In taxa of the subfamiliesPereskioideae andOpuntioideae the stomata are generally oriented parallel to the stem axis with only minor deviations. The stomata of taxa of the subfamilyCactoideae do in general not show a particular orientation with the notable exception of a number of epiphytic genera from the tribeHylocereeae.—The results of these investigations in general coincide well with the generic classification of theCactaceae byHunt (1967). A few cases where stomatal characters suggest a differing classification merit further investigations.—Additionally, the possibilities to distinguish between paracytic and parallelocytic stomata are discussed and an amended definition for the latter is given.     

Androecial development and systematics inFlacourtiaceae s. l.

Androecial development of 13 species belonging to six tribes ofFlacourtiaceae has been investigated. While inScolopieae andFlacourtieae the stamens develop centrifugally, inErythrospermeae, Oncobeae andPangieae they are initiated in a centripetal sequence or a sequence that is neither distinctly centripetal nor centrifugal. The distribution of these developmental patterns coincides with the distribution of other characters (e.g. cyanogenic compounds, salicoid leaf teeth) and therefore supports a split of the family intoFlacourtiaceae s. str. (containing theScolopieae, Homalieae, Prockieae, Flacourtieae, Casearieae andBembicieae) andKiggelariaceae (withErythrospermeae, Oncobeae andPangieae) and is in accordance with results of recentrbcL studies.     

rbcL sequences support exclusion ofRetzia,Desfontainia, andNicodemia from theGentianales

The taxonomic positions ofRetzia, Desfontainia, andNicodemia have been much discussed, and all three genera have been included inLoganiaceae (Gentianales). We have made a cladistic analysis ofrbcL gene sequences to determine the relationships of these taxa toGentianales. Four newrbcL sequences are presented; i.e., ofRetzia, Desfontainia, Diervilla (Caprifoliaceae), andEuthystachys (Stilbaceae). Our results show thatRetzia, Desfontainia, andNicodemia are not closely related toLoganiaceae or theGentianales. Retzia is most closely related toEuthystachys and is better included inStilbaceae. The positions ofDesfontainia andNicodemia are not settled, butDesfontainia shows affinity for theDipsacales s.l. andNicodemia for theLamiales s.l.     

Chloroplast DNA restriction site variation in theVernonieae (Asteraceae), an initial appraisal of the relationship of New and Old World taxa and the monophyly ofVernonia

Chloroplast DNA restriction site variation was examined for 35 taxa in theVernonieae and four outgroup tribes, using 17 restriction enzymes mapped for ca. 900 restriction sites per species; 139 mutations were found to be phylogenetically informative. Phylogenetic trees were constructed using Wagner and weighted parsimony, and evaluated by bootstrap and decay analyses. Relationships of Old and New World taxa indicate complex geographical relationships; there was no clear geographic separation by hemisphere. The relationships between Old and New World Vernonias found here support prior morphological analyses. The sister group to all New and most Old World taxa was composed of a small group of Old World species including yellow-flowered, trinervate-leaved species previously postulated to be basal in the tribe. The majority of both New and Old World taxa are derived from a lineage beginning with the monotypic genusStokesia, an endemic of the southeastern United States. The genusVernonia was also found to be paraphyletic within both the New and Old World. Available data do not support either the separation ofVernonia or the tribeVernonieae into geographically distinct lineages. The pattern of relationships within theVernonieae for taxa from North America, Asia, Africa, Central and South America is most similar to that of several other groups of both plants and animals with a boreotropical origin, rather than an origin in Gondwanaland. Such a pattern of distribution suggests more ancient vicariant events than are routinely postulated for theAsteraceae.     

The phylogeny and evolution of thePterygodium—Corycium complex (Coryciinae,Orchidaceae)

The leaf-anatomy, palynology, seed-morphology, vegetative morphology and especially the highly complicated floral morphology of theCoryciinae s. str. (Diseae: Orchidoideae: Orchidaceae) are described and illustrated in detail. On the basis of these characters the presumed phylogeny, based on a rigorous cladistic analysis, is presented. The cladistic biogeographical analysis of theCoryciinae s. str. shows that it is a member of the Afrotemperate Track, with a pattern of vicariance events typical of the members of this track. An analysis of the patterns of speciation shows that allopatric speciation appears to be rare, and that parapatric speciation across edaphic boundaries may be the most important factor. Proceeding from the information presented, a new classification of the group is proposed in which we recognize the four generaCeratandra, Evotella, Pterygodium andCorycium. The new monotypic genusEvotella comprises a species originally described asPterygodium rubiginosum. The three species of the genusAnochilus are transferred toCorycium andPterygodium. P. magnum, which was originally described asPterygodium but was transferred toCorycium lately, is placed in a monotypic section ofPterygodium.     

Flavonoid distribution of Mulinum (Apiaceae, Hydrocotyloideae, Mulineae)

Flavonoid chemistry of the genusMulinum and selected Mulineae taxa was studied. Both flavones and flavonols were identified as C- and O-glycosides. AllMulinum species contain 6,8-di-C-glycosyl chrysoeriol (flavone) and, with the exception of one, quercetin (flavonol). The presence of both flavones and flavonols in this genus weakens previous generalizations that the mulineae contain only flavonols and are primitive compared to other Apiaceae tribes. Based on the selected taxa studied,Azorella appears to differ from bothMulinum andGymnophyton in producing more kinds of flavonols, andGymnophyton appears similar toMulinum in the production of both chrysoeriol and quercetin as well as relatively few compounds. The flavonoid profile ofAsteriscium glaucum is reported as well. In general, a more homogeneous flavonoid compound composition for the Apiaceae is suggested.     

Phylogenetic relationships in the Hamamelidaceae: Evidence from the nucleotide sequences of the plastid genematK

The Hamamelidaceae is a family that bridges the basal elements of the Rosidae and the lower Hamamelidae, thus a better understanding of the phylogeny of the family is important for clarifying evolutionary patterns in the diversification of eudicots. However, subfamilial as well as tribal relationships in the Hamamelidaceae have been controversial. Nucleotide sequences of the chloroplast genematK were used to study the intergeneric relationships of the family. In the phylogenetic trees, constructed using parsimony analysis, the clade containingAltingia andLiquidambar (Altingioideae) is sister to a clade that includes all other Hamamelidaceae.Exbucklandia andRhodoleia form a clade, suggesting a close relationship between the two genera.Disanthus is sister to the monophyletic Hamamelidoideae. The paraphyletic arrangement ofDisanthus, Mytilaria andExbucklandia with respect to the Hamamelidoideae does not support the combination of these genera in one subfamily. In the Hamamelidoideae, thematK phylogeny supports the monophyly of several previously recognized groups with modifications, including the tribes Eustigmateae (incl.Molinadendron), Fothergilleae (excl.Molinadendron andMatudaea), and the subtribe Dicoryphinae. However, the Hamamelideae as traditionally circumscribed is polyphyletic. Apetaly has evolved three times independently in the Hamamelidoideae.     

Phylogenetic relationships of epacrids and vaccinioids (Ericaceae s. l.) based onmatK sequence data

Cladistic relationships of epacrids and vaccinioids (Ericaceae) are investigated using nucleotide sequence data from the chloroplast encodedmatK gene. Sequences of 56 taxa were aligned and analyzed using parsimony methods. Results show thatVaccinioideae as currently recognized are not monophyletic. The epacrids are sister to a clade that includes theLyonia group, theGaultheria group, and theVaccinieae. Arbutus andPyrola branch early inEricaceae, before the rhododendroid group.Enkianthus is sister to the remainingEricaceae (includingEpacridaceae).Vaccinieae are strongly supported as monophyletic, butVaccinium andAgapetes are polyphyletic.