Plagiochila virginica A.Evans rather than P. dubia Lindenb. & Gottsche occurs in Macaronesia; placement in sect. Contiguae Carl is supported by ITS sequences of nuclear ribosomal DNA

 Plagiochila dubia Lindenb. & Gottsche is reduced to a synonym of the Neotropical P. patula (Sw.) Lindenb. Specimens from the Canary Islands and Madeira proved to belong to the eastern North American P. virginica A.Evans, new to Europe. Phylogenetic analyses of the internal transcribed spacer regions (ITS1-5.8S-ITS2) of nuclear ribosomal DNA of ten Plagiochila species produced four independent lineages that are well supported by all bootstrap analyses (maximum likelihood, maximum parsimony, and distance). These lineages correspond with the Plagiochila sections Arrectae, Contiguae, Cucullatae and Glaucescentes. Spruce's “Ramiflorae” and “Cauliflorae” may no longer be regarded as monophyletic units of Plagiochila. Received August 19, 2001 Accepted October 11, 2001     

Systematics ofPlagiochila sect.Glaucescentes Carl (Hepaticae) from tropical America: A morphological and chemotaxonomical approach

The systematics of theGlaucescentes Carl, a poorly known neotropical section of the large genusPlagiochila, is revised based on a large set of gametophytic, sporophytic and chemical characters. Three species are recognized,P. buchtiniana Steph.,P. longispina Lindenb. and Gottsche andP. diversifolia Lindenb. and Gottsche, the latter with two chemotypes. Twelve binomina are reduced to synonymy. The species occur in montane regions of tropical America, especially in the northern and central Andes. In addition,P. longispina is newly reported from the Azores, where it was previously known asP. allorgei Herzog and Perss. TheGlaucescentes are excellently characterized by morphological and chemical features and are clearly separated from theContiguae Carl, a group often confused with theGlaucescentes. Morphologically, theGlaucescentes stand apart by the brown oil bodies, thinwalled inner capsule wall cells and partly unequally bispiral elaters. Chemically, they are characterized by the accumulation of partially hydrogenated bibenzylderivatives (longispinone, longispinol), 3-benzylphthalides and various flavonoids, and by the low production of terpenoids.Publication No. 142 of the Arbeitskreis Chemie und Biologie der Moose, Universität des Saarlandes, Saarbrücken.     

Surface wax, a new taxonomic feature in Plagiochilaceae

A scanning electron microscope study of 81 species of Plagiochilaceae revealed the presence of superficial waxes on the leaves and stems ofPlagiochilion mayebarae and 5 species ofPlagiochila. The waxes are not visible in the light microscope and were unknown in Plagiochilaceae.Plagiochila fuscolutea andP. longiramea (=P. sect.Fuscoluteae) are characterised by the predominant occurrence of membraneous wax platelets;Plagiochila aerea, P. rudischusteri andP. tabinensis(=P. sect.Bursatae) predominately form various types of wax rodlets. Our findings show for the first time the systematic usefulness of leaf surface waxes in the liverworts.P. tabinensis contains surface waxes in amounts of ca. 1.4% dry weight composing of steryl esters, triacylglycerols and free fatty acids.     

nrDNA internal transcribed spacer data reveal that Rhodoplagiochila R.M. Schust. (Marchantiophyta: Jungermanniales) is a member of Plagiochila sect. Arrectae Carl

The monospecific genus Rhodoplagiochila, previously known only from the type collected in the Venezuelan Andes, has been collected at a second site close to the type locality. Up to now, generic status of Rhodoplagiochila has been accepted by most authors, but the taxon had been alternately assigned to Plagiochilaceae or Lophoziaceae. Phylogenetic analyses of nrITS sequences from Rhodoplagiochila and other representatives of the Plagiochilaceae clearly reveal the former to be in a clade with members of the Neotropical–Atlantic European Plagiochila sect. Arrectae. Within the Arrectae, Rhodoplagiochila is positioned in a robust clade made up of several accessions of Plagiochila bifaria. Morphologically, Rhodoplagiochila differs from other phenotypes of P. bifaria by the papillose leaf surface and a leaf margin which is ciliately toothed all-around. As an outcome of the molecular and morphological investigations, Rhodoplagiochila rosea is treated as a variety of P. bifaria, and the new combination P. bifaria var. rosea comb. et stat. nov. is proposed. In tropical America the family Plagiochilaceae seems to be represented only by Plagiochila.     

Sesquiterpenoids of fourteen Plagiochila species

The distribution of ent-2,3-secoaromadendrane-, ent-aromadendrane-, ent-bicyclogermacrane- and ent-maaliane-type sesquiterpenoids in fourteen Plagiochila species is described. These sesquiterpenes are the significant chemosystematic markers of Plagiochila. The intense pungent substance of some Plagiochila species is due to an ent-2,3-secoaromadendrane-type sesquiterpene hemiacetal, plagiochiline A.     

13C NMR chemical shifts of ovalifoliene and related compounds with the 2,3-seco-alloaromadendrane skeleton: Structure of (+)-9α-acetoxyovalifoliene,a plant growth inhibitor

¹³C NMR spectra of several 2,3-seco-alloaromadendrane-type acetyl hemiacetals were analysed and the structure of an additional plant growth-inhibitor was determined to be (+)-9α-acetoxyovalifoliene. The biological activity of the new compound is also described.     

Reinstatement of Lophocoleaceae (Jungermanniopsida) based on chloroplast gene rbcL data: exploring the importance of female involucres for the systematics of Jungermanniales

Maximum likelihood analysis of 113 rbcL sequences leads to a well resolved phylogeny of Jungermanniales. All species with perigynia or marsupia are found in one clade, whereas species with coelocaules are placed in several lineages. The broadly circumscribed Geocalycaceae (including Lophocoleaceae) of most recent authors are resolved as polyphyletic. Geocalycaceae genera which develop female involucres without involvement of stem tissue (Chiloscyphus, Heteroscyphus, Leptoscyphus, Physotheca) form a robust clade which is placed sister to Plagiochilaceae whereas the genera with involucres originating at least partly from stem tissue (Geocalycaceae s.str., Geocalyx, Harpanthus, Saccogyna) are nested within the paraphyletic Jungermanniaceae. This topology leads to the exclusion of the strictly perianth-bearing species from Geocalycaceae and the reinstatement of Lophocoleaceae. Campanocolea is nested within Chiloscyphus. Physotheca and Chiloscyphus breutelii are placed within an unsupported clade with several accessions of Leptoscyphus. Heteroscyphus forms a paraphyletic grade at the base of Chiloscyphus.     

Biogeography of Plagiochila (Hepaticae): natural species groups span several floristic kingdoms

Aim This paper presents a synthesis of our recent results regarding the biogeography of Plagiochila using a molecular approach, and documents intercontinental ranges within this largest genus of the hepatics. Methods A maximum likelihood analysis of sixty‐one nrITS sequences of Plagiochila was performed and the molecular topology obtained was compared with morphological, phytochemical and geographical data. Results Our molecular data set allowed the identification of eleven Plagiochila sections, the majority of which cover at least two floristic kingdoms. Seven sections have species in Europe (sect. Arrectae, Carringtoniae, Fuscoluteae, Glaucescentes, Plagiochila, Rutilantes, Vagae). Plagiochila species from Atlantic Europe are usually close to or conspecific with neotropical taxa, whereas species widespread in Europe are closely related to Asian ones and not to those in the Neotropics. Plagiochila sect. Arrectae represents a neotropical – Atlantic European clade. The section is not closely related – as has often been suggested – to the morphologically similar sect. Zonatae from Asia and western North America. Sequence data show that the African P. integerrima and the neotropical P. subplana are members of the Asian sect. Cucullatae (sect. Ciliatae, syn. nov.), which becomes pantropical in distribution. An ITS sequence of P. boryana from Uganda confirms the Afro‐American range of the primarily neotropical sect. Hylacoetes. Similarities in sporophyte morphology between the sect. Cucullatae and sect. Hylacoetes are the result of parallel evolution. Main conclusions Our results indicate that intercontinental ranges at section and species level are common in Plagiochila. Carl's (1931) subdivision of Plagiochila into sections restricted to one floristic kingdom is outdated. Biogeographical patterns in Plagiochila are not dissimilar to those of other groups of bryophytes but elucidation of the geographical ranges of the taxa requires a molecular approach. Contrary to earlier belief, most Plagiochila species from Atlantic Europe do not have close relatives in Asia but are conspecific with or closely related to species from tropical America.     

Plagiochila cucullifolia var. anomala var. nov. from Ecuador, with notes on discordant molecular and morphological variation in Plagiochila

Plagiochila cucullifolia Jack & Steph. var. anomala J. Heinrichs & Gradst. var. nov. is described and illustrated. The new variety is known from a single locality in southern Ecuador and differs from P. cucullifolia var. cucullifolia by the flat, not saccate leaves, somewhat smaller plant size and weaker leaf dentation. According to phylogenetic analyses of 35 nrDNA ITS1 and ITS2 sequences of Plagiochila, the two varieties of P. cucullifolia form a monophyletic lineage and are placed in a well supported clade together with five other species of Plagiochila sect. Hylacoetes: P. dimorpha Lindenb. & Gottsche var. ecuadorica (Inoue) J. Heinrichs, P. flabelliflora Steph., P. patriciae J. Heinrichs & H. Anton, P. macrostachya Lindenb. and P. turgida Herzog. Within Plagiochila, nrITS sequence variation is not concordant with morphological diversification. ITS sequences of Plagiochila cucullifolia s.str. and of P. dimorpha var. ecuadorica differ in only 23 aligned positions whereas two sequences of P. subplana Lindenb. differ in 86 aligned positions. Morphologically, P. cucullifolia s.str. and P. dimorpha var. ecuadorica differ in more than 20 characters and previously these two taxa were placed in separate genera. P. subplana phenotypes show considerable variation in leaf shape and dentation but the extremes are linked by numerous intermediates.