Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological andndhF sequence data

Phylogenetic relationships within the tribe Antirrhineae (Scrophulariaceae) are analysed and discussed on the basis of parsimony analyses of morphological andndhF gene sequence data. The results indicate that the tribe Antirrhineae consists of four major groups of genera, theAnarrhinum clade, theGambelia clade, theMaurandya clade, and theAntirrhinum clade. TheAnarrhinum clade, consisting of the Old World bee-pollinated generaAnarrhinum andKickxia, is sister to the rest of the tribe. TheGambelia clade consists of the New World generaGambelia andGalvezia, which are very closely related and pollinated by hummingbirds. TheMaurandya clade consists of one subclade includingMaurandya and a number of related bee- or hummingbird-pollinated New World genera and another subclade with the Old World bee-pollinated generaAsarina andCymbalaria. TheAntirrhinum clade consists mainly of bee-pollinated Old World genera, such asAntirrhinum, Linaria, Chaenorhinum, and their segregates, but also includes the New World generaMohavea andHowelliella, of which the latter is known to be partly pollinated by hummingbirds. It is concluded that hummingbirdpollination has evolved independently within Antirrhineae at least three times from bee-pollinated ancestors.     

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.     

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.     

Phylogeny and biosystematics ofPseudomonotes (Dipterocarpaceae) based on molecular and morphological data

The placement of a recently discovered South American monotypic genus,Pseudomonotes tropenbosii, in subfam.Monotoideae (Dipterocarpaceae) extends the geographical range of the subfamily from Africa to the Neotropics. Although morphological and anatomical evidence suggest similarities betweenPseudomonotes andMonotes, the close alliance of these two genera was questionable due to their disjunct distribution and a lack of phylogenetic analysis. In the present study, we reconstructed the phylogeny ofPseudomonotes and other putatively related taxa usingrbcL sequence data. The analysis ofrbcL sequences of 20 taxa belonging to 15 genera and eight families recovered a single most parsimonious tree. The genusSarcolaena (Sarcolaenaceae) formed a clade sister to the monophyleticDipterocarpaceae clade.Monotes andPseudomonotes formed a strongly supported group, sister to the monophyletic clade withPakaraimaea and the remaining Asiatic dipterocarp species studied. The study strongly supports the placement ofPseudomonotes within subfam.Monotoideae of theDipterocarpaceae.     

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.     

New classification of liverworts based on molecular and morphological data

Complete sequences for the 18S-rRNA gene of 22 bryophytes (12 completely new) were determined and used to construct phylogenetic trees. The evaluation of sequence data according to the maximum parsimony principle (PAUP 3.1.1) and the neighbor-joining method (MEGA) results in similar phylogenetic trees in which theBryopsida appear as a sister group to theJungermanniopsida, and both together as a sister group to theMarchantiopsida. Among theMarchantiopsida, theSphaerocarpales diverge early as a separate clade. TheMetzgeriales andJungermanniales are monophyletic. They belong to one clade and cannot be separated by either method of evaluation.     

Elektronenmikroskopische Untersuchungen zur Frage der verwandtschaftlichen Beziehungen zwischenTheligonum undRubiaceae: Feinbau der Siebelement-Plastiden und Anmerkungen zur Struktur der Pollenexine

Theligonum cynocrambe and 13 species ofRubiaceae contain S-type sieve-element plastids, wide-spread in Dicotyledons. Alignment ofTheligonum toCaryophyllales (Centrospermae), especiallyPhytolaccaceae, is unlikely, because this order is characterized by specific P-type plastids. SEM investigations show the pollen exine ofTheligonum to be microreticulate, with additional supratectate spinules.Asperula and other genera of the tribeRubieae have a tectum perforatum (punctitegillate sexine), also with supratectate verrucae or spinulae.—Thus ultrastructure suggests (but not definitely proves) relationships betweenTheligonum andRubiaceae, while affinities betweenTheligonum andCaryophyllales are excluded.

     

A phylogenetic analysis of the palm subtribe Oncospermatinae (Arecaceae) based on morphological characters

Subtribe Oncospermatinae (Arecaceae: Arecoideae: Areceae) is a diverse group of spiny Old World palms. The subtribe includesOncosperma, a widespread Asian genus of five species, along with seven monotypic genera, all endemic to the Seychelles and Mascarene Islands of the western Indian Ocean. A phylogenetic analysis was conducted in order to test the monophyly of subtribe Oncospermatinae with respect to other Old World genera of tribe Areceae. A matrix of 38 morphological characters was scored for 29 taxa, including 11 species of the Oncospermatinae. A single most parsimonious tree was found, resolving the subtribe as a polyphyletic group of two distinct clades. One clade containingAcanthophoenix, Deckenia, Oncosperma, andTectiphiala was placed as sister to a large group that includes members of subtribes Archontophoenicinae, Arecinae, Iguanurinae, and Ptychospermatinae. The other clade of Oncospermatinae, including the Seychelles endemic generaNephrosperma, Phoenicophorium, Roscheria, andVerschaffeltia, was resolved as sister to the Madagascar endemic subtribe Masoalinae, and may have arisen in the western Indian Ocean region.     

Systematics and karyoevolution inMagnoliidae:Tetrameranthus as compared with otherAnnonaceae genera of the same chromosome number

First generic chromosome counts reveal the base number x=7 for the generaTetrameranthus andRollinia. T. umbellatus from the Peruvian Amazon is diploid (2n=14),T. duckei from Brazil (Manaus) is tetraploid (2n=28). In the NeotropicsRollinia (7 species counted) has developed diploid to octoploid taxa (2n=14, 28, 42, 56). Counts of 7 South AmericanAnnona species are presented for comparison (2n=14, 28). The West AfricanCleistopholis patens has 2n=14. The Asian genusMezettia: 2n=14 and the neotropicalGuatteria tribe: 2n=28 are also revised. A detailed karyomorphological comparison, including karyotypes, banding patterns, condensing behaviour of chromosomes and structure of interphase nuclei reveals that the closely related generaAnnona andRollinia are almost identical in their diploid genomes, whereas the polyploid ones differ in their heterochromatin (=hc) composition and number of NO-chromosomes.Cleistopholis, Mezettia and theGuatteria tribe are karyologically and systematically distinct from each other and fromAnnona/Rollinia. Tetrameranthus as compared with the karyomorphology of about 60 other Annonaceous genera has a very peculiar and unusual karyomorphology which underlines its isolated position. Nuclear structures are almost identical in the African genusUvariopsis (2n = 16) and partly similar in theGuatteria tribe; both also share some morphological similarities and possibly are related. From a comparison ofTetrameranthus with several nuclear types within theMagnoliidae, a new model of chromosome evolution in primitive Angiosperms is suggested. In respect to their eco-morphological differentiation the genera investigated differ strongly from each other.Dedicated to Prof. Dr. K.-H.Rechinger on the occasion of His 80th birthday.     

Characterization and phylogenetic distribution of a chloroplast DNA rearrangement in theBerberidaceae

Restriction site maps and a clone bank of chloroplast DNA (cpDNA) ofMahonia higginsae (Munz)Ahrendt (Berberidaceae) were constructed. The size ofMahonia cpDNA was about 167 kb. Precise mapping using gene probes revealed that cpDNA ofM. higginsae has an inverted repeat (IR) 11.5 kb larger than the tobacco IR. The expansion of the IR into the large single copy region has resulted in the duplication of at least ten genes includingpsbB. The phylogenetic distribution of the expanded IR was examined in twenty-five species ofBerberis andMahonia, twenty species representing the fifteen remaining genera of theBerberidaceae, and four species from four allied families. Our survey indicates that only the species of the closely related generaBerberis andMahonia share the 11.5kb expansion of IR. This result supports their close phylogenetic relationship, which has been suggested previously by chromosomal, morphological, and serological data.     

Phylogenetic relationships among genera in theLiliaceae-Asparagoideae-Polygonatae s.l. inferred fromrbcL gene sequence data

The chloroplast gene encoding ribulose-1,5-bisphosphate-carboxylase (rbcL) was sequenced for phylogenetic analysis of 13 species (10 genera) in the tribePolygonatae s.l. of theLiliaceae-Asparagoideae. The data were analysed using maximum parsimony and neighbour-joining methods. There were 233 phylogenetically informative sites out of 1368 base pairs compared. The results suggest that there are three monophyletic groups withinPolygonatae s.l. with high bootstrap confidence values. Group A representsPolygonatae s.str., with generaMaianthemum, Smilacina, Convallaria, Disporopsis, andPolygonatum. Group B containsUvularia andDisporum and group C includesStreptopus, Tricyrtis, Clintonia, andProsartes. The study suggests thatPolygonatae s.l. are not a monophyletic group, including at least three groups of different phylogenetic origin. Monophyly of the taxa within groups A, B, and C is supported by the high bootstrap confidence values (85–100%) of the bootstrap replications for both parsimony and neighbour-joining methods. The differences between each group (calculated as 100x base substitutions per site) were 6.99–9.03 for group A and B, 4.92–7.35 for A and C, and 6.66–7.57 for B and C.     

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.     

Karyosystematic studies onRubiaceae: Chromosome counts from Sri Lanka

Chromosome numbers for 16 different taxa ofRubiaceae from Sri Lanka are presented. Counts for 13 species, including 5 genera and 1 tribe, are new. The results are listed, compared with available literature data and discussed in regard to their systematical importance.     

Relationships, circumscription, and biogeography of Arcytophyllum (Rubiaceae) based on evidence from cpDNA

A cladistic analysis was performed using nucleotide sequence variation in therps16 intron and thetrmL-F region (plastid DNA).Arcytophyllum belongs in a subclade of the tribe Spermacoceae (s.l.) together with the American species presently classified in the generaHedyotis andHoustonia. This subclade is morphologically characterized by cymbiform seeds.Arcytophyllum is the sister group of all AmericanHedyotis andHoustonia and it is suggeste that these latter would be most conveniently treated as a single genus, the correct name of which would beHoustonia.Arcytophyllum should be circumscribed such that it excludesA. serpyllaceum, which is not a member of theArcytophyllum-Houstonia clade but more closely related toBouvardia. The phylogeny that was reconstructed suggests that the ancestral area of theArcytophyllum-Houstonia clade is the South American tectonic plate.     

Evolution of flower shape inVeroniceae (Scrophulariaceae)

Floral evolution in the tribeVeroniceae was examined using phylogenetic analysis combining 24 adult morphology and chromosome number characters with 22 qualitative and quantitative floral development characters. Taxa sampled included nine species ofVeroniceae and as an outgroup one species each ofDigitaleae andVerbasceae. Veronica, Besseya, andSynthyris formed one clade, subtended byPseudolysimachion and then by theHebe group;Veronicastrum orWulfenia represent the basal-most branch of the tribe. The ancestral flowers of theVeroniceae may have been small with moderately short corolla tubes and lobes; long corolla tubes arose four times in the tribe and large corolla lobes twice.     

Ribes (Grossulariaceae) phylogeny as indicated by restriction-site polymorphisms of PCR-amplified chloroplast DNA

We surveyed exemplars from all 12 infrageneric taxa ofRibes (Grossulariaceae) for restriction site variation in two cpDNA regions, fromrbcL toaccD and fromrpoC1 torpoC2, in order to develop an explicit phylogenetic hypothesis and to assess the validity of infrageneric classifications. Maximum parsimony analysis resolves sect.Ribes (red currants), sect.Berisia (European alpine currants), sect.Symphocalyx (golden currants), sect.Grossularia plus sect.Grossularioides (true gooseberries and spiny currants), andHesperia, Lobbia, and probably sect.Robsonia (west North American gooseberries) as well-supported monophyletic groups. The clade of sectionsGrossularioides andGrossularia is unexpected, and suggests that subgenusGrossularia is not monophyletic. Alternatively, sect.Grossularioides may have acquired its cpDNA via hybridization and introgression. SectionsCoreosma (black currants) andHeritiera (dwarf currants) are apparently non monophyletic. Relationships among the well-supported lineages and the other sampled taxa remain unresolved. Maximum likelihood analysis is consistent with the parsimony results.     

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.     

Fruit differentiation, palynology, and systematics in theScabiosa group of genera andPseudoscabiosa (Dipsacaceae)

Fruits ofDipsacaceae are single-seeded, have bristle-shaped calyx segments and are tightly enclosed by four fused bracts forming an epicalyx. Comparative morphological and anatomical studies reveal a great diversity of epicalyx and calyx, often relevant to fruit dispersal. The present contribution deals with theScabiosa group of genera, the core of theScabioseae tribe. Most of its taxa develop a diaphragma from a meristem on the inside of the epicalyx. This diaphragma, together with the lower part of the epicalyx encloses the fruit proper, whereas the upper parts form a so-called epi-diaphragma (ed) and a ± hyaline corona. Differences of the epicalyx with respect to the size and position of the ed, elaboration of the corona, origin of pits (=foveoles) and other morphological and anatomical specializations can be demonstrated. Together with palynological and karyological data these new facts support an improved concept of relationships and systematics for the taxa studied:Scabiosa sect.Scabiosa and sect.Cyrtostemma are closely related and should be united to form the genusScabiosa s. str.;Pycnocomon can be maintained as an independent genus, sister toScabiosa sect.Trochocephalus which then has to be treated as a genus,Lomelosia. In contrast, the following genera have to be included inLomelosia:Tremastelma asLomelosia sect.Callistemma, andScabiosiopsis as part ofLomelosia sect.Lomelosia. Pseudoscabiosa deviates in so many features that it has to be excluded from the redefinedScabioseae s. str.