Phylogenetic relationships ofSphaerophoraceae (Ascomycetes) inferred from SSU rDNA sequences

SSU rDNA was sequenced from the lichenized fungiBunodophoron scrobiculatum andLeifidium tenerum (Sphaerophoraceae), andStereocaulon ramulosum andPilophorus acicularis (Stereocaulaceae) and analysed by maximum parsimony with 44 homologous ascomycete sequences in a cladistic study. A small insertion (c. 60 nt.) was found in the sequence ofLeifidium tenerum. Sphaerophoraceae constitutes a strongly supported monophyletic group which groups together withLecanora dispersa and theStereocaulaceae. Together withPorpidia crustulata, this larger group is a sistergroup to thePeltigerineae. This analysis thus supports theLecanorales as monophyletic, includingSphaerophoraceae and thePeltigerineae, but does not provide strong support for this monophyly. The analysis also suggests that the prototunicate ascus in theSphaerophoraceae is a reversion to the plesiomorphic state. Based on morphological, anatomical and chemical reasons,Sphaerophoraceae is proposed to belong to one of the groups presently included in the paraphyletic suborderCladoniineae within theLecanorales.     

The familial and subfamilial relationships ofApocynaceae andAsclepiadaceae evaluated withrbcL data

Sequence data for therbcL gene from twenty-four taxa of the familiesApocynaceae andAsclepiadaceae were cladistically analysed in order to evaluate the existing familial and subfamilial classification. The taxa sampled represent all described subfamilies and a majority of the described tribes. The cladistic analysis shows that theAsclepiadaceae are nested within theApocynaceae. An amalgamation of the two families is therefore recommended. The subfamilial classification is also in need of revision: the subfamiliesPlumerioideae andApocynoideae of the current classifications are paraphyletic, as are many of the tribes. Potential subfamily candidates and characters traditionally used in the classification are discussed.     

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 cladistic analysis ofAnisopappus (Asteraceae: Inuleae)

A cladistic analysis of the genusAnisopappus (Asteraceae: Inuleae) has been undertaken. A hypothesis of species interrelationships in the genus is presented for the first time. The analysis also includedArctotis (Arctoteae), used as outgroup, and five additional genera from theInuleae: Geigeria, Calostephane, Asteriscus, Buphthalmum, Pulicaria, andInula. It is concluded thatAnisopappus is a monophyletic group situated at the base of the tribe, diagnosed by, e.g., their obtuse stylar sweeping-hairs. The species with acute sweeping-hairs were found to be derived within the genus. Problems concerning species delimitation, biogeography and character evolution in the genus are briefly discussed.     

Heliantheae sensu lato (Asteraceae), clades and classification

The interrelationships within theHeliantheae s. lato and the closely relatedEupatorieae are analyzed and discussed. The basis to this discussion is a cladistic analysis of 141 morphological characters (172 apomorphic states) scored for 97 genera. TheHeleniae s. lato, a subgroup of theHeliantheae s. lato, are paraphyletic, and a monophyletic group corresponding largely to theHeliantheae s. str. is recognized. TheEcliptinae sensuRobinson are polyphyletic. TheCoreopsidinae form an ingroup in theHeliantheae s. str. TheTageteae (Pectidinae) and theMadieae (Madiinae) are two separate branches within the helenioid assemblage of taxa.     

Cladistic interrelationships and major clades of theEricales

A cladistic analysis of theEricales based on morphological, anatomical, embryological, and phytochemical data has been undertaken. Resulting from the analysis, 56 equally parsimonious cladograms were obtained. The tribeEnkiantheae was found to be the sister-group of the rest of theEricaceae, in which theEmpetraceae, theEpacridaceae, Pyrolaceae, andMonotropaceae are forming derived ingroups. It is thus concluded that theEricaceae are a paraphyletic assemblage as presently circumscribed, and thatEmpetraceae andEpacridaceae are better treated as members of theEricaceae.     

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.     

P-type sieve-element plastids and the systematics of theArales (sensuCronquist 1988)—with S-type plastids inPistia

The sieve-element plastids of 126 species of theArales were investigated by transmission electron microscopy. With the exception ofPistia (with S-type plastids) all contained the monocotyledon specific subtype-P2 plastids characterized by cuneate protein crystals. While the species studied from bothAcoraceae andLemnaceae have form-P2c plastids (i.e., with cuneate crystals only), those of theAraceae belong to either form P2c (14 species), P2cs (the great majority) or P2cfs (Monstera deliciosa, only, with form-P2cs plastids in the otherMonstera species studied). The form-P2cs plastids of theAraceae are grouped into different categories according to the quantity and quality of their protein and starch contents. The subfamilyLasioideae is redefined to comprise all aroid P2c-taxa and those P2cs-genera that contain only one or very few starch grains. Only little starch is also recorded in the sieve-element plastids ofGymnostachys (Gymnostachydoideae), with the other plastid data denying a close relationship toAcorus. While equal amounts of starch and protein are generally present in sieve-element plastids of the subfamiliesPothoideae, Monsteroideae, Colocasioideae, Philodendroideae, andAroideae, maximum starch content and only very few protein crystals are found in form-P2cs plastids ofCalla (Calloideae),Ariopsis (Aroideae), andRemusatia (Colocasioideae?). In the latter, both morphology and size of sieve-element plastids are close to those ofPistia.—In theAraceae the diameters of the sieve-element plastids exhibit a great size range, but are consistent within a species and within a defined part of the plant body. Comparative data are mainly available for stem and petiole sieve-element plastids.—The accumulated data are used to suggest an affiliation of the species to subfamilies and to discuss the phylogeny of theArales. Forms and sizes of their plastids support a separation of bothAcoraceae andLemnaceae from theAraceae. The presence of S-type plastids inPistia does not favour direct and close relationships to the form-P2c genusLemna.—The prevailing form-P2cs plastids might support proposals that place theArales (together with also form-P2cs plastid containingDioscoreales) in the neighbourhood of basal dicotyledons. BesidesAsarum andSaruma (Aristolochiaceae), with monocotyledonous form-P2c plastids,Pistia (with dicotyledonous S-type plastids) gives another example for a link between the two angiosperm classes.     

Phylogeny of theAsterales sensu lato based onrbcL sequences with particular reference to theGoodeniaceae

TherbcL gene of 25 taxa was sequenced and analyzed cladistically in order to define more precisely the orderAsterales s.l. and to reconstruct the phylogeny ofGoodeniaceae. The cladistic analyses show that theAsterales comprise the familiesAbrophyllaceae, Alseuosmiaceae, Argophyllaceae, Asteraceae, Calyceraceae, Campanulaceae s.l.,Donatiaceae, Goodeniaceae (includingBrunoniaceae),Menyanthaceae, Pentaphragmataceae, andStylidiaceae. Abrophyllaceae, Alseuosmiaceae, Brunoniaceae, andDonatiaceae have previously not been studied in this respect. Within theGoodeniaceae, four groups supported by therbcL data can be distinguished: the genusLechenaultia, theAnthotium-Dampiera-group, the genusBrunonia, and a group formed by the remaining genera, theScaevola-Goodenia-group.     

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.     

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.     

Phylogeny of theCyclanthaceae

The affinities ofCyclanthaceae are discussed, and it is concluded that the sister-group to this family is most probablyPandanaceae. A hypothesis of generic relationships inCyclanthaceae, based on cladistic methods, is presented. Bootstrap analysis and Bremer support (decay index) have been used to test the strength of individual clades, and the result is compared with previously made phylogenetic analyses. TheSphaeradenia group (Chorigyne, Stelestylis, Sphaeradenia, andLudovia) is supported as monophyletic and acceptably resolved, while theAsplundia group (remaining genera inCarludovicoideae) may be paraphyletic, with largely uncertain relationships. A formal recognition of these groups is therefore not justified. The probable character evolution inCarludovicoideae is discussed.     

Pollen grain morphology ofCoris (Primulaceae)

Pollen grain morphology, sculpturing, and wall ultrastructure are investigated in two species ofCoris (Primulaceae),C. monspeliensis L. andC. hispanica Lange. The study includes both acetolysed and unacetolysed pollen. No evidence of any major palynological difference is recorded between these two species, apart from a somewhat larger pollen inC. monspeliensis. However,Coris can be distinguished from the remaining members of thePrimulaceae by the conjunction of relatively large pollen grains, prominent margo, and particular tectal pattern causing a reticulate surface with minute luminal perforations decreasing towards the colpi. From both these distinctive features, and others typically primulaceous, some evolutionary considerations are inferred. Finally, the higher proportion of irregular grains inC. hispanica is interpreted in light of environmental stress.     

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.     

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.     

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.     

Chloroplast DNA variation and phylogeny of theRanunculaceae

Restriction site mapping of chloroplast DNA from 31 species representing 26 genera of theRanunculaceae was performed using eleven restriction endonucleases. The chloroplast genome varies in length from approximately 152 to 160 kb. Length variants are frequent in theRanunculaceae and range from usually less than 300 bp to rarely 1.5 kb. The inverted repeat is extended into the large single copy (LSC) region by 4–4.5 kb inAnemone, Clematis, Clematopsis, Hepatica, Knowltonia, andPulsatilla. Several inversions are present in the LSC-region of the cpDNA in all these genera and inAdonis. The frequency of restriction site mutations varies within the chloroplast genome in theRanunculaceae between 4 and 32 mutations per kilobase, and is lowest in the inverted repeat and the regions containing the ATPase-genes and the genespsaA, psaB, psbA, rpoB, andrbcL. A total of 547 phylogenetically informative restriction sites was utilized in cladistic analyses of the family using Wagner, Dollo, and weighted parsimony. These three parsimony analyses result in different tree topologies. Four, six, and one equally most parsimonious trees were obtained with Wagner, Dollo, and weighted parsimony, respectively. The amount of support for the monophyletic groups was evaluated using bootstrapping and decay analysis. All three parsimony methods suggest thatHydrastis is the sister group to the remainder of theRanunculaceae, and that theAnemone-Clematis group, which shares several derived cpDNA rearrangements, is monophyletic. Only a few of the traditional groups in theRanunculaceae are supported by cpDNA restriction side data. Only Dollo parsimony provides support for the hypothesis thatThalictroideae andRanunculoideae are monophyletic.