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—A systematic revision of the tropical moss genus Papillaria was carried out by combining morphological studies with phylogenetic reconstructions based on molecular data from the plastid (trnL-F and rpl16) and the nuclear (ITS region) genomes. For the morphological study a set of qualitative and quantitative features was studied in over 2800 herbarium specimens, including the types, of all 55 previously accepted species in the genus. After revision, nine species of Papillaria were recognized, Papillaria crocea and P. flexicaulis which are broadly distributed in the tropics, P. africana that is endemic to Africa, P. laevifolia that is known from the Americas, and P. flavolimbata, P. funiformis, P. leuconeura, P. nitens, and P. zeloflexicaulis that are known from Australia and the Pacific Islands. The majority of the other previously accepted names do not belong to Papillaria s. s. but to the closely related genera Toloxis and Meteorium. We propose five new synonyms under P. africana as well as one new synonym for P. crocea and one for P. laevifolia. The revision reduced the number of Papillaria species in Africa from ten to one and in South America from thirteen to three. We extend the geographic range of P. crocea due to a new record from eastern South America (Brazil).

We present a description of the new species Rhynchostegium occultum from central Chile, including illustrations, a distribution map, and preliminary molecular data that supports its taxonomic placement. The new taxon is restricted to coastal relict forests of central Chile. An overview of the Chilean taxa of Rhynchostegium, and a key for local species are also provided. The new combination Rhynchostegium corralense, and new synonyms for Rhynchostegium acanthophyllum and for Cratoneuropsis chilensis are proposed. Oxyrrhynchium hians is newly reported for Chile, apparently being a recent introduction. Lectotypes are selected for Hypnum corralense, Hypnum acanthophyllum, and for Rhynchostegiella acanthophylla var. robusta.

Centro de Investigación Forestal (CIFOR), Nonrecombining sex chromosomes, like the mammalian Y, often lose genes and accumulate transposable ele ments, a process termed degeneration. The correlation between suppressed recombination and degeneration is clear in animal XY systems, but the absence of recombination is confounded with other asymmetries between the X and Y. In contrast, UV sex chromosomes, like those found in bryophytes, experience symmetrical population genetic conditions. Here, we generate nearly gapless female and male chromosome-scale reference genomes of the moss Ceratodon purpureus to test for degeneration in the bryophyte UV sex chromosomes. We show that the moss sex chromosomes evolved over 300 million years ago and expanded via two chromosomal fusions. Although the sex chromosomes exhibit weaker purifying selection than autosomes, we find that suppressed recombination alone is insufficient to drive degeneration. Instead, the U and V sex chromosomes harbor thousands of broadly expressed genes, including numerous key regulators of sexual development across land plants., This work was supported by NSF DEB-1541005 and 1542609 and start-up funds from UF to S.F.M.; microMORPH Cross-Disciplinary Training Grant, Sigma-Xi Grant-In-Aid of Research, and Society for the Study of Evolution Rosemary Grant Award to S.B.C.; NSF DEB-1239992 to N.J.W.; the Emil Aaltonen Foundation and the University of Turku to S.O.; and NSF DEB-1541506 to J.G.B. and S.F.M. The work conducted by the U.S. Department of Energy Joint Genome Institute was supported by the Office of Science of the U.S. Department of Energy under contract no. DE-AC02-05CH11231., 12 Pág. Supplementary material for this article is available at http://advances.sciencemag.org/cgi/ content/full/7/27/eabh2488/DC1

Non-recombining sex chromosomes, like the mammalian Y, often lose genes and accumulate transposable elements, a process termed degeneration1,2. The correlation between suppressed recombination and degeneration is clear in animal XY systems1,2, but the absence of recombination is confounded with other asymmetries between the X and Y. In contrast, UV sex chromosomes, like those found in bryophytes, experience symmetrical population genetic conditions3,4. Here we test for degeneration in the bryophyte UV sex chromosome system through genomic comparisons with new female and male chromosome-scale reference genomes of the moss Ceratodon purpureus. We show that the moss sex chromosomes evolved over 300 million years ago and expanded via two chromosomal fusions. Although the sex chromosomes show signs of weaker purifying selection than autosomes, we find suppressed recombination alone is insufficient to drive gene loss on sex-specific chromosomes. Instead, the U and V sex chromosomes harbor thousands of broadly-expressed genes, including numerous key regulators of sexual development across land plants.

Phylogenetic position of Homalothecium laevisetum and relationship with the genus Palamocladium

In recent treatments of Fissidens subgenus Fissidens, the species delimitation of F. bryoides, the type species of the subgenus, has been rather variable. In Finland, F. bryoides has either been split into F. bryoides, F. gymnandrus and F. viridulus, or treated as a single species, F. bryoides s. lato, including two or more intraspecific taxa. The herbarium material of Finnish F. bryoides, F. gymnandrus and F. viridulus was studied to clarify the distribution and ecological requirements of the three taxa and to find out which morphological and ecological characters are best for their delimitation. The phylogeny lends support to recognizing F. bryoides s. lato including F. gymnandrus and F. viridulus as F. bryoides var. gymnandrus and F. bryoides var. viridulus. The phylogeny reconstruction based on 18 morphological characters that are commonly used to distinguish the taxa could not resolve F. bryoides, F. gymnandrus and F. viridulus as monophyletic species. However, F. bryoides s. lato including all herbarium material of the three species was resolved as a monophyletic group morphologically distinct from the closely related F. pusillus. The synapomorphic characters that distinguish F. bryoides s. lato are the percurrent costa, limbidia that reach the leaf acumen, the acuminate leaf acumen, and a preference for substrates other than rock. We found differences in habitat requirements between the varieties, while sexual systems sometimes vary even within a single specimen. Fissidens bryoides var. bryoides and F. bryoides var. gymnandrus appear to have wider distribution in Finland than expected. Unlike F. bryoides var. viridulus the number of localities of F. bryoides var. bryoides and F. bryoides var. gymnandrus is low. Information on their conservation status is also provided.

Mosses (division Bryophyta) are characterized by the dominance of haploid, poikilohydric gametophytes, and relatively persistent sporophytes that are dependent on the gametophyte generation. Becaus...

— The distribution and systematic position of Kindbergia Ochyra in China is reevaluated. The phylogenetic analyses based on nuclear marker (ITS1-5.8S-ITS2) and chloroplastic markers (rpl16, trnG, and trnL-F) show that the alleged species of “Kindbergia” in China and Himalayas form a maximally supported clade (1.00 PP and 100 MPBS), which is closely related to the clade of Brachythecium Schimp., Eurhynchiadelphus Ignatov et Huttunen, and Myuroclada Besch., whereas the other Kindbergia specimens from Africa, America, Europe, Japan and the Southern Hemisphere form a monophyletic group, which is sister to Scleropodium Schimp. The specimens of Chinese species of “Kindbergia” are found, both molecularly and morphologically, to be identical with the Himalayan K. dumosa (Mitt.) Ignatov et Huttunen, and differ from Kindbergia specimens from other regions in the world in the shape of proximal branch leaves, operculum shape and capsule orientation. A new monotypic genus Pseudokindbergia M. Li, Y.F. Wang, Ig...

In the light of a phylogenetic reconstruction based on a multi-loci data set including sequence data from all three compartments (plastome: (rps4)–trnF, rpl16 group II intron; chondrome: nad5 group I intron; nuclear: ITS1&2) the Orthostichellaceae are described as a new family of pleurocarpous mosses. It is distributed mainly in the Neotropics and tropical Africa, and provisionally contains the genera Dixonia (Asia, 1 species), Afrothamnium gen. nov. (Africa and Neotropics, 1 sp.), Pinnatidendron gen. nov. (Africa and Neotropics, 1 sp.), Deslooveria gen. nov. (Africa and French Guiana, 5 spp.), Scabrellifolium gen. nov. (Neotropics and Africa, 2 spp.), and Orthostichella (Neotropics and Africa, 9 spp.). In the Neckeraceae we describe Dannorrisia gen. nov. (N. America, 1 sp.), Austrothamnium gen. nov. (Australasia, 2 spp.), and Bryobuckia gen. nov. (endemic to Chile, 7 spp.). Porotrichum s.str. becomes an essentially Neotropical genus and it contains the formerly recognized genera Porotrichopsis and Porotrichodendron. Thamnobryum s.str. is distributed exclusively in the Northern Hemisphere and is largely temperate. The mainly Macaronesian Echinodiaceae becomes a family of two genera, Echinodium (3 spp.) and Pseudomalia gen. nov. (1 sp.). The morphological evolution in the Orthostichellaceae is discussed based on the phylogeny.

The systematic position of the genus Bryhnia Kaurin has been problematic and remains unsettled. A phylogenetic analysis of Bryhnia and closely related genera, based on the nuclear marker ITS1–5·8S–ITS2 and the plastid markers rpl16, trnG, and trnL–F, was carried out to clarify their relationships and systematic positions. The phylogenetic trees generated from Maximum parsimony and Bayesian analyses show that Bryhnia s.s., represented by B. novae-angliae (Sull. & Lesq.) Grout, B. hultenii E.B.Bartram, and B. scabrida (Lindb.) Kaurin, forms a maximally supported clade nested within Brachythecium Schimp. ‘Bryhnia scabrida’ sensu Ignatov & Huttunen (2002) is Bryhnia brachycladula Cardot, and together they form a monophyletic group with Eurhynchiadelphus eustegia (Besch.) Ignatov & Huttunen and Myuroclada maximowiczii (G.G.Borshch.) Steere & W.B.Schofield. We propose the inclusion of the species currently recognized in Bryhnia in Brachythecium, with the transfer of Bryhnia brachycladula to the genus Myuroclada...

The phylogenetic position and generic composition of the moss family Plagiotheciaceae were explored using DNA sequence data from three genomes: plastid trnL-F and rps4, mitochondrial nad5 intron and nuclear ITS1-5.8S-ITS2. Our phylogenetic analyses included 35 terminals from Plagiotheciaceae and 71 outgroup taxa from a representative set of hypnalean moss families. The family Plagiotheciaceae is resolved in the early-diverging Hypnales grade, together with Fontinalaceae, Habrodontaceae and several genera which are mainly distributed in the area of the former Gondwanan supercontinent. However, monophyly of the family can only be attained if the three Southern Hemisphere genera, Acrocladium, Catagonium and Rhizofabronia, are excluded. Ancestral state reconstruction for eight morphological characters reveals that many characters used to delimit the family, such as a lack of pseudoparaphyllia and rhizoids inserted in the leaf axils, were already present in the ancestor of Hypnales. Dispersal–vicariance analysis suggests that Plagiotheciaceae and Fontinalaceae have their ancestral distributions in the area of the former Laurasian supercontinent. As the analyses also reveal a Gondwanan distribution for the ancestor of Hypnales in general, Plagiotheciaceae and Fontinalaceae represent the first diverging Laurasian lineages in the order. © 2013 The Linnean Society of London

The antimicrobial activity and phenolic compounds of five Finnish honey products against important human pathogens Streptococcus pneumoniae, S. pyogenes, Staphylococcus aureus, and methicillin-resistant S. aureus were analyzed. Microbroth dilution method and HPLC-DAD were used in antimicrobial testing and phenolic compound determination, respectively. Significant antimicrobial activity (p

The pleurocarpous moss genus Forsstroemia (Neckeraceae) that was taxonomically revised in 1987 on morphological grounds is now revisited using sequence data from two plastid regions (rps4-trnF and rpl16) and nuclear ribosomal DNA (ITS1 and 2). The taxon sampling comprises 11 out of 13 species and a representative set of Neckeraceae and Lembophyllaceae taxa as outgroups. Forsstroemia japonica was placed next to Shevockia inunctocarpa in a clade only remotely related to Forsstroemia. Thus, F. japonica is transferred to Shevockia. Forsstroemia remotifolia is renamed as Leptodon remotifolius and Pseudopterobryum tenuicuspis as Forsstroemia tenuicuspis (syn. F. tripinnata). The nine species in Forsstroemia form a monophyletic group, six of the species being restricted to Asia, one (F. coronata) to South America, and two (F. producta and F. trichomitria) having a wider distribution. A previously suggested hybrid origin for F. coronata was not supported by the sequence data.

The Hypnales are the largest order of mosses comprising approximately 4200 species. Phylogenetic reconstruction within the group has proven to be difficult due to rapid radiation at an early stage of evolution and, consequently, relationships among clades have remained poorly resolved. We compiled data from four sequence regions, namely, nuclear ITS1–5.8S–ITS2, plastid trnL–F and rps4, and mitochondrial nad5, for 122 hypnalean species and 34 species from closely related groups. Tree topologies from both Bayesian and parsimony analyses resolve the order as monophyletic. Although inferences were made from fastevolving genes, and despite strong phylogenetic signal in the nuclear ITS1–5.8S–ITS2 data, monophyly, as well as backbone nodes within the Hypnales, remains rather poorly supported except under Bayesian inferences. Ancestral distribution based on Bayesian dispersal-vicariance analysis supports a Gondwanan origin of the Hypnales and subsequent geographical radiation in the area of the former Laurasian supercontinent. Reconstruction of historical biogeography is congruent with mainly tropical and Gondwanan distributions in the sister groups Hypnodendrales, Ptychomniales, and Hookeriales, and with the dating for the oldest pleurocarp and hypnalean fossils. We contrast groupings in the phylogenetic tree with recent classifications and other phylogenetic inferences based on molecular data, and summarise current knowledge on the evolutionary history of, and relationships among, the Hypnales.

Bryophyte floras typically exhibit extremely low levels of endemism. The interpretation, that this might reflect taxonomic shortcomings, is tested here for the Macaronesian flora, using the moss species complex of Rhynchostegium riparioides as a model. The deep polyphyly of R. riparioides across its distribution range reveals active differentiation that better corresponds to geographic than morphological differences. Morphometric analyses are, in fact, blurred by a size gradient that accounts for 80% of the variation observed among gametophytic traits. The lack of endemic diversification observed in R. riparioides in Macaronesia weakens the idea that the low rates of endemism observed in the Macaronesian bryophyte flora might solely be explained by taxonomic shortcomings. To the reverse, the striking polyphyly of North American and European lineages of R. riparioides suggests that the similarity between the floras of these continents has been over-emphasized. Discriminant analyses point to the existence of morphological discontinuities among the lineages resolved by the molecular phylogeny. The global rate of error associated to species identification based on morphology (0.23) indicates, however, that intergradation of shape and size characters among species in the group challenges their identification.

Bacterial adhesion to the cell surface is a crucial step before infection can take place. Inhibition of bacterial binding offers a novel preventive approach against infections. Cranberry (Vaccinium macrocarpon Ait.) juice has been found to have antiadhesive activity against different bacteria. Streptococcus pneumoniae is an important pathogen and the most common cause for pneumonia, meningitis, and otitis media. In this study the inhibitory activity of cranberry (Vaccinium oxycoccos L.), bilberry (Vaccinium myrtillus L.) and crowberry (Empetrum nigrum and Empetrum hermaphroditum L.) juice fractions against pneumococcal binding was tested using human bronchial cells (Calu-3) as an adhesion model. In addition, the antimicrobial activity of the berry juice fractions was tested. It was found that the studied berry juice fractions had antiadhesion activity and cranberry juice was the most active. The adhesion inhibition activity of cranberry juice was nearly 90% at a concentration of 8.7 mg/g of soluble solids. The antimicrobial activity of the studied berry juice fractions was found to be remarkable; pneumococcal growth was inhibited totally at a concentration of ∼86 mg/g. Both antiadhesion and antimicrobial activities were reduced after solid-phase extraction of the berry juices, which may suggest molecular synergistic effects of the berry juice molecules against S. pneumoniae. The findings indicate that cranberry, bilberry and crowberry juices have potential against pneumococcal infections.

The adhesion of pathogens to host tissues is the requirement for the initiation of the majority of infectious diseases. It was shown recently that the binding of Neisseria meningitidis pili to immobilized human epithelial cells is inhibited by molecular size fractions (10-100 kDa) of berry juices. Additionally, the isolated meningococcal pili bound to polyphenolic fractions of berry juices. The present study investigated the antiadhesive effects of berry juice polyphenolics against living meningococcal bacteria in a human epithelial cell culture model. The ability of bilberry, cranberry, crowberry and lingonberry juice polyphenolic fractions to inhibit the attachment of N. meningitidis bacteria to HEC-1B human epithelial cells in a cell culture model was examined. The antibacterial effect of the fractions was tested using a microtiter broth microdilution assay. The most effective adhesion inhibition of 75% was achieved with cranberry juice polyphenolic fraction followed by crowberry (63%), bilberry (63%) and lingonberry (57%) juice polyphenolic fractions. Bacterial survival rates after incubation with the fractions varied between 75-100%. The present results suggest berry juice polyphenols as inhibitors of adherence of N. meningitidis. Thus the binding of meningococci to berry juice polyphenols might be protective for the host against the infection.

Phylogenetic relationships of the aquatic moss genus Platyhypnidium and terrestrial species of Rhynchostegium were explored using a phylogeny based on three DNA sequence regions, the nuclear ITS1-5.8S-1TS2, and the plastid trnL-F and trnD-T. This is the first study using trnD-T in the phyogeny of mosses. Platyhypnidium appeared polyphyletic and most of its species were found within a Rhynchostegium clade including species from four different genera: Rhynchostegium, Eriodon, Eurhynchiella and Platyhypnidium. Use of the name Rhynchostegium requires conservation against Eriodon. Within Rhynchostegium s.l., aquatic Rhynchostegium species (i.e., Platyhypnidium) were resolved in four geographically delimited groups: an Afro-European group, an American―East Asiatic group, a Malesian-Australian group and a group comprising the South American P. fuegianum together with Eriodon and Eurhynchiella. Topologies from independent analyses ofplastid and nuclear data did not show conflict, and the occurrence of hybridzation events in the clade was therefore not supported. The phylogeny indicates numerous habitat shifts between terrestrial and aquatic habitats, as well as between different terrestrial (epiphytic and epigeic) habitats, which may have affected taxonomic complexity in Rhynchostegium. New morphological characters, such as a striate leaf lamina, axillary hairs composed of relatively short cells, and clearly differentiated cells of the costa, separate Platyhypnidium species within Eurhynchioideae from those in the Oxyrrhyuchium-Donrichardsia clade (Helicodontioideae). Two morphologically unique specimens, Platyhypnidium muellerii from Hawaii and Platyhypnidium sp. from Northern Australia, are described as Donrichardsia bartramii and Rhynchostegium brevinerve. The new genus Hedenaesia (Helicodontioideae) is established to accommodate Platyhypnidium austrinum.

Fine-scale spatial genetic structure (SGS) of the liverwort, Barbilophozia attenuata, occupying an area characterized by a network of ant trails, was investigated using microsatellite markers. This is the first study investigating SGS in a liverwort. Significant genetic differentiation was detected among colonies along and outside ant trails, and the SGS pattern varied depending on the spatial scale. At short distances, kinship coefficients were significantly positive up to about eight meters, after which they approached zero and turned negative, while at distances greater than 25 m the values were about zero. Thus, nearby individuals are more closely related than expected, at mid-distances less related, and at great distances genotypes show a random distribution. We suggest that the reproductive mode strongly affects SGS in B. attenuata. Asexual propagation by relatively large gemmae allows more effective establishment than sexual reproduction by small-sized spores, and causes an aggregation of similar genotypes, although the inbreeding effect cannot be ruled out. In environments with small-scale disturbances, e.g., ant trails, gemmae are favoured over spores at establishment. Also, the diaspore bank of the forest floor can be activated by disturbances, which may affect SGS. At mid-distances, the isolation by distance effect, presumably related to comparatively ineffective gemma dispersal, is visible, while at greater distances, the role of spores as effective means of dispersal is evident. The Sp statistic values, which quantify the strength of SGS, indicate that outsider colonies possess less SGS than do plant colonies along ant trails, which relates to the more frequent spore production of outsider colonies. Moreover, dispersal from fallen logs or stumps may be more effective than dispersal from ground-level colonies along ant trails. Apparently, ants do not have much role as dispersal vectors, nor do the physical structures of ant trails as dispersal corridors, although they provide open areas for colonization.

Rhynchostegium confusum, a new species from Spain and the Iberian Peninsula is described. It is closely related to R. confertum, from which it differs in its noticeably wider stem leaves with wider and apically spurred costa, more flexuous branch leaves, and shorter seta. It also differs in its habit and habitat: the plants are tightly attached to siliceous rocks just over the water level in rivulets that dry out during the summer in the Mediterranean region. Molecular data show that the new species derives from R. confertum, but is clearly differentiated from the studied populations of the latter in a number of missing haplotypes.

The Macaronesian endemic flora has traditionally been interpreted as a relict of a subtropical element that spanned across Europe in the Tertiary. This hypothesis is revisited in the moss subfamily Helicodontioideae based on molecular divergence estimates derived from two independent calibration techniques either employing fossil evidence or using an Monte Carlo Markov Chain (MCMC) to sample absolute rates of nucleotide substitution from a prior distribution encompassing a wide range of rates documented across land plants. Both analyses suggest that the monotypic Madeiran endemic genus Hedenasiastrum diverged of other Helicodontioideae about 40 million years, that is, well before Macaronesian archipelagos actually emerged, in agreement with the relict hypothesis. Hedenasiastrum is characterized by a plesiomorphic morphology, which is suggestive of a complete morphological stasis over 40 million years. Macaronesian endemic Rhynchostegiella species, whose polyphyletic origin involves multiple colonization events, evolved much more recently, and yet accumulated many more morphological novelties than H. percurrens. The Macaronesian moss flora thus appears as a complex mix of ancient relicts and more recently dispersed, fast-evolving taxa.

The moss genus Rhynchostegiella (Helicodontioideae, Brachytheciaceae) has long served as a convenient repository for small brachythecioid pleurocarps. Its circumscription is revised in the context of a chloroplast phylogeny of the Helicodontioideae employing trnL-trnF, atpB-rbcL, psbT-psbH, and psbA-trnH sequence data. The analysis resolves with full posterior probabilities a core Rhynchostegiella clade of eight species. Rhynchostegiella pumila and R. duriaei are both resolved outside that clade and accommodated in their own genera, Microeurhynchium gen. nov. and Pseudorhynchostegiella gen. nov., respectively. Rhynchostegiella leptoneura is sister to Aerolindigia capillacea and R. papuensis is closely related to Eurhynchiella zeyheri. One of the reasons why these unrelated species, together with other taxa, were traditionally included within Rhynchostegiella, is that the genus is morphologically poorly defined by only a single synapomorphic change followed by reversals in half of the species. The ...

Antiadhesion therapy is a promising approach to the fight against pathogens. Antibiotic resistance and the lack of effective vaccines have increased the search for new methods to prevent infectious diseases. Previous studies have shown the antiadhesion activity of juice from cultivated cranberries (Vaccinium macrocarpon Ait.) against bacteria, especially E. coli. In this study, the binding of two streptococcal strains, Streptococcus pneumoniae and Streptococcus agalactiae, to molecular size fractions (FI, FII and FIII,10 kDa, 10-100 kDa, and100 kDa, respectively) of berries and berry and fruit juices from 12 plant species were studied using a microtiter well assay. For Streptococcus suis a hemagglutination inhibition assay was used. In general, binding activity was detected especially to wild cranberry (Vaccinium oxycoccos L.) and to other Vaccinium species. S. pneumoniae cells bound most to cranberry juice fraction FI and S. agalactiae cells to cranberry fraction FIII. Hemagglutination induced by S. suis was most effectively inhibited by cranberry fraction FII. NMR spectra of some characteristic active and non-active fractions were also measured. They indicate that fractions FII and FIII contained proanthocyanidins and/or other phenolic compounds. The results suggest Vaccinium berries as possible sources of antiadhesives against bacterial infections.

Although the Lembophyllaceae has undergone considerable revision during the last century, the generic and familial level relationships of this pleurocarpous moss family are still poorly understood. To address this problem, a generic revision of the Lembophyllaceae based on molecular data was undertaken. We analyzed two plastid markers, the trnL-trnF and the psbT-psbH region in combination with the ITS2 of nuclear ribosomal DNA. The molecular data reveal that the current circumscription of the family is too narrow and that several genera previously placed in the Lembophyllaceae should be reincluded. The family includes: Bestia, Camptochaete, Dolichomitra, Dolichomitriopsis, Fallaciella, Fifea, Isothecium, Lembophyllum, Looseria stat. nov., Pilotrichella, Rigodium, Tripterocladium, and Weymouthia. Looseria contains a single species: Looseria orbiculata comb. nov. Acrocladium is excluded and provisionally accommodated in the Lepyrodontaceae. Generic limits supported by the molecular data support a return to the early twentieth century family concept of Brotherus. The analyses indicate that the segregate genus Orthostichella is distinct from its parent genus Pilotrichella, probably at the family level. Whereas Pilotrichella is resolved within the Lembophyllaceae, Orthostichella clusters with Porotrichum and Porothamnium forming a clade (OPP-clade) sister to the remaining Neckeraceae and Lembophyllaceae. Hence, the Neckeraceae is paraphyletic. Recognition of the OPP-clade as a new family is desirable but awaits the results of detailed ongoing morphological studies.

Blocking bacterial adhesion to host surfaces provides novel potential to control infections. The present study was directed to binding and inhibitory activity of different fresh berries and berry and fruit juices against Neisseria meningitidis . Berries and juices were fractionated according to their molecular size into three fractions. A microtiter well assay for binding of N. meningitidis pili to berry and juice fractions was constructed. In addition, adhesion inhibition to human epithelial cells (HEC-1B) was tested. The active fractions were then subfractionated by employing solid-phase extraction. Subfractions were characterized by RP-HPLC-DAD, and the pili binding was evaluated by using microtiter well binding assay. Binding and inhibitory activity were detected to bilberry, cranberry, lingonberry, and crowberry fractions, which contained anthocyanins or a mixture of proanthocyanidins and flavonols. Thus, the findings identify several previously unknown binding and inhibitory activities and may suggest Vaccinium berries and crowberry as promising sources against meningococcal adherence.

Competing hypotheses that rely either on a stepping-stone dispersal via the North Atlantic or the Bering land bridges, or more recent transoceanic dispersal, have been proposed to explain the disjunct distribution of Mediterranean flora in southern Europe and western North America. These hypotheses were tested with molecular dating using a phylogeny of the moss genus Homalothecium based on ITS, atpB-rbcL, and rpl16 sequence data. The monophyly of two main lineages in Western Palearctic (Europe, central Asia and north Africa) and North America is consistent with the ancient vicariance hypothesis. The monophyly of Madeiran H. sericeum accessions supports the recognition of the Macaronesian endemic H. mandonii. A range of absolute rates of molecular evolution documented in land plants was used as probabilistic calibration prior by a Bayesian inference implementing a relaxed-clock model to derive ages for the nodes of interest. Our age estimates for the divergence of the American and Western Palearctic Homalothecium clade (5.7 Ma, IC 3.52-8.26) and the origin of H. mandonii (2.52 Myr IC 0.86-8.25) are not compatible with the ancient vicariance hypothesis. Age estimates suggests that species distributions result from rare instances of dispersal and subsequent sympatric diversification. The calibrated phylogeny indicates that Homalothecium has undergone a fast radiation during the last 4 Myr, which is consistent with the low levels of morphological divergence among sibling species.

Results from our analysis of chloroplast trnL-trnF and nuclear ITS1 and ITS2 sequence data and 35 morphological characters for 134 taxa of pleurocarpous mosses provide an evidence for resurrecting two moss families, Pseudoleskeaceae and Pylaisiaceae. Both these families were described by Schimper in 1860, but soon afterwards included in Leskeaceae and Hypnaceae, respectively, and apparently never used in the 20 t h century. However, sequence-level data analysis (chloroplast encoded trnL intron and trnL-trnF spacer, nuclear encoded ITS1 & ITS2), and combined molecular and morphological analysis demonstrate the remote position of (1) the Lescuraea-group from the main part of Leskeaceae; and (2) a group of species belonging to Pylaisia, Homomallium and Hypnum (sections others than sect. Hypnum} from Hypnum sect. Hypnum (with H. cupressiforme as the type of the genus).

Brachytheciaceae is often considered a taxonomically difficult group of mosses. For example, morphological variation has led to difficulty in generic delimitation. We used DNAsequence data (chloroplast psbT-H and trnL-F and nuclear ITS2) together with morphology (63 characters) to examine the relationships within this family. The combined unaligned length of the DNA sequences used in the phylogenetic analyses varied between 1277 and 1343 bp. For phylogeny reconstruction we performed direct optimization, as implemented in POY. Analyses were performed with three different gap costs and the morphological data partition was weighted both: (1) equal to gap cost, and (2) with a weight of one. The utility of sensitivity analysis has recently been cast into doubt; hence in this study it was performed only to explore the effects of weighting on homology statements and topologies and to enable more detailed comparisons between earlier studies utilizing the direct optimization method. The wide sequence length variation of non-coding ITS2 sequences resulted in character optimizations (i.e., ‘‘alignments’’) of very different lengths when various gap costs were applied. Despite this variation, the topologies of equally parsimonious trees remained fairly stable. The inclusion of several outgroups, instead of only one, was observed to increase the congruence between data sets and to slightly increase the resolution. An inversion event in the 9 bp loop region in the chloroplast psbT-N spacer in mosses has been postulated to include only uninformative variation, thus possibly negatively impacting the phylogeny reconstruction. Despite this inversion, its variation within Brachytheciaceae was clearly congruent with information from other sources, but inclusion of these 9 bp in the analysis had only a minor effect on the phylogenetic results. In the most parsimonious topology, which was obtained with equal weighting of all data, Meteoriaceae and Brachytheciaceae were resolved as monophyletic sister groups, which had recently been suggested based on a few shared morphological characters. Our study revealed some new generic relationships within the Brachytheciaceae, which are discussed in light of the morphological characters traditionally used for generic delimitation. � The Willi Hennig Society 2004.

The tandemly arranged genes psbT, psbN, and psbH code for proteins of photosystem II and are located in the large single copy region (LSC) of the chloroplast genome, downstream of psbB. So far, most of the studies dealing with this region have been interested in the organization and transcription of the psbB operon, while less is known about the transcription of psbN or the phylogenetic utility of this region. In the current study we discuss a sigma70-type bacterial promoter motif upstream of psbN and present its consensus sequence for bryophytes. An analysis of the 3′ flanking inverted repeat sequences revealed a dyad symmetrical element, which is able to form a stable stem-loop structure. This hairpin structure is characterised for land plants, with an emphasis on bryophytes. Furthermore, we observed an inversion of up to 9 bases in the loop region of the hairpin structure, which occasionally occurred in different unrelated bryophyte families, orders and classes. Small inversions are frequently obscured in the alignment since, during automatic alignment, many gap-weighting schemes may not introduce gaps, nor does a manual insertion of gaps always seem needed at first glance. In subsequent phylogenetic analyses, minute inversion may overweight a particular mutation by interpreting the single inversion event as multiple apomorphic substitutions, which is particularly problematic since such inversion events are known to often be highly homoplastic. In the present study we use the psbT-N spacer as an example to quantify the effect of such small inversions, contrasting the phylogenetic structure obtained with and without information from the hairpin loop. We show that obscured minute inversions can highly significantly reduce the robustness of the phylogenetic hypothesis inferred from the data set.

The reproduction of Pohlia nutans (Hedw.) Lindb. and Pleurozium schreberi (Brid.) Mitt. were compared in the surroundings of copper smelters at Harjavalta, Finland. Samples of both species were collected from 16 sample plots, which were situated on a 12-km transect at intervals of half a kilometre from the smelters. As a result of pollution the production of gametangia decreased in P. schreberi, and near the smelters most of the shoots were sterile. The biomass of shoots seems to have an effect on the production of perichaetia; reproducing females were always the largest ones in the colonies. The reproductive strategy of Pohlia nutans also varied. While, in moderately clean areas, colonies reproduced sexually and formed sporophytes, in the vicinity of the smelters caducous shoot apices were produced. The numbers of sporophytes in colonies were highest in moderately polluted areas and decreased further away from the smelters. The numbers of spores per capsule in P. nutans decreased at polluted site...

Two closely related tropical genera from the pleurocarpous moss family Neckeraceae are revised: the second largest genus in the family, Neckeropsis , currently with 29 species, and Himantocladium , comprising six species. Twenty-one species of Neckeropsis and five of Himantocladium were included in this study, which is based on phylogenetic analyses using sequence level data from the plastid ( rps4 ) -trnT-trnL-trnF cluster and rpl16 as well as nuclear ITS1 & 2. Neckeropsis appeared as polyphyletic. Neckeropsis s. str. comprises 12 species and a further four species, not included in the analysis, are tentatively retained in the genus based on morphology. Four new genera are segregated from Neckeropsis : Pengchengwua (with one sp.), Planicladium (two spp.), Pseudoparaphysanthus (five spp.), and Neckeromnion (five spp.). Neckeropsis gracilis is synonymized with Neckeromnion urocladum and reported for the first time from Laos. Four molecular analyzed species are resolved in the Himantocladium s. str. clade, with a fifth species being added solely on morphological basis. Himantocladium formosicum is transferred to Neckeropsis . Leaf-like paraphyses (ramenta), unique in Neckeropsis s. str. , have evolved two times. In addition, the rare Noguchiodendron sphaerocarpum from the Himalayan region, the single species in its genus, was included in a phylogenetic analysis for the first time and it was found to be closely related to the Asian genus Taiwanobryum .

Adaptive evolution has often been proposed to explain correlations between habitats and certain phenotypes. In mosses, a high frequency of species with specialized sporophytic traits in exposed or epiphytic habitats was, already 100 years ago, suggested as due to adaptation. We tested this hypothesis by contrasting phylogenetic and morphological data from two moss families, Neckeraceae and Lembophyllaceae, both of which show parallel shifts to a specialized morphology and to exposed epiphytic or epilithic habitats. Phylogeny-based tests for correlated evolution revealed that evolution of four sporophytic traits is correlated with a habitat shift. For three of them, evolutionary rates of dual character-state changes suggest that habitat shifts appear prior to changes in morphology. This suggests that they could have evolved as adaptations to new habitats. Regarding the fourth correlated trait the specialized morphology had already evolved before the habitat shift. In addition, several other specialized “epiphytic” traits show no correlation with a habitat shift. Besides adaptive diversification, other processes thus also affect the match between phenotype and environment. Several potential factors such as complex genetic and developmental pathways yielding the same phenotypes, differences in strength of selection, or constraints in phenotypic evolution may lead to an inability of phylogeny-based comparative methods to detect potential adaptations.

The family Neckeraceae is composed of three distinct clades, of which two, i.e. the Neckera and Thamnobryum clades, are well defined. The third clade, consisting of species belonging to Caduciella, Curvicladium, Handeliobryum, Himantocladium, Homaliodendron, Hydrocryphaea, Neckera, Neckeropsis, Pinnatella, Shevockia and Taiwanobryum, is the focus of this study. Based on sequence data from the trnS-rps4-trnT-trnL-trnF plastid cluster and the rpl16 intron as well as from nuclear ITS1&2, the phylogenetic relationships of these genera are reconstructed. The nearest relatives of this clade are resolved shedding more light on the evolution of the family. The generic composition of the clade and its individual genera are discussed; polyphyly requires redefinition of Pinnatella, Neckeropsis and Homaliodendron. The positions of Touwia and Homalia within the family are addressed in an additional analysis based on more extensive sequence data, and the corresponding new combinations are made. Several further taxonomic changes are proposed, including Circulifolium gen. nov., comprising the former Homaliodendron exiguum and H. microdendron.

Earlier phylogenetic studies, including species belonging to the Neckeraceae, have indicated that this pleurocarpous moss family shares a strongly supported sister group relationship with the Lembophyllaceae, but the family delimitation of the former needs adjustment. To test the monophyly of the Neckeraceae, as well as to redefine the family circumscription and to pinpoint its phylogenetic position in a larger context, a phylogenetic study based on molecular data was carried out. Sequence data were compiled, combining data from all three genomes: nuclear ITS1 and 2, plastid trnS-rps4-trnT-trnL-trnF and rpl16, and mitochondrial nad5 intron. The Neckeraceae have sometimes been divided into the two families, Neckeraceae and Thamnobryaceae, a division rejected here. Both parsimony and Bayesian analyses of molecular data revealed that the family concept of the Neckeraceae needs several further adjustments, such as the exclusion of some individual species and smaller genera as well as the inclusion of the Leptodontaceae. Within the family three well-supported clades (A, B and C) can be distinguished. Members of clade A are mainly non-Asiatic and nontropical. Most species have a weak costa and immersed capsules with reduced peristomes (mainly Neckera spp.) and the teeth at the leaf margins are usually unicellular. Clade B members are also mainly non-Asiatic. They are typically fairly robust, distinctly stipilate, having a single, at least relatively strong costa, long setae (capsules exserted), and the peristomes are well developed or only somewhat reduced. Members of clade C are essentially Asiatic and tropical. The species of this clade usually have a strong costa and a long seta, the seta often being mammillose in its upper part. The peristome types in this clade are mixed, since both reduced and unreduced types are found. Several neckeraceous genera that were recognised on a morphological basis are polyphyletic (e.g. Neckera, Homalia, Thamnobryum, Porotrichum). Ancestral state reconstructions revealed that currently used diagnostic traits, such as the leaf asymmetry and costa strength are highly homoplastic. Similarly, the reconstructions revealed that the 'reduced' sporophyte features have evolved independently in each of the three clades. Earlier phylogenetic studies, including species belonging to the Neckeraceae, have indicated that this pleurocarpous moss family shares a strongly supported sister group relationship with the Lembophyllaceae, but the family delimitation of the former needs adjustment. To test the monophyly of the Neckeraceae, as well as to redefine the family circumscription and to pinpoint its phylogenetic position in a larger context, a phylogenetic study based on molecular data was carried out. Sequence data were compiled, combining data from all three genomes: nuclear ITS1 and 2, plastid trnS-rps4-trnT-trnL-trnF and rpl16, and mitochondrial nad5 intron. The Neckeraceae have sometimes been divided into the two families, Neckeraceae and Thamnobryaceae, a division rejected here. Both parsimony and Bayesian analyses of molecular data revealed that the family concept of the Neckeraceae needs several further adjustments, such as the exclusion of some individual species and smaller genera as well as the inclusion of the Leptodontaceae. Within the family three well-supported clades (A, B and C) can be distinguished. Members of clade A are mainly non-Asiatic and nontropical. Most species have a weak costa and immersed capsules with reduced peristomes (mainly Neckera spp.) and the teeth at the leaf margins are usually unicellular. Clade B members are also mainly non-Asiatic. They are typically fairly robust, distinctly stipilate, having a single, at least relatively strong costa, long setae (capsules exserted), and the peristomes are well developed or only somewhat reduced. Members of clade C are essentially Asiatic and tropical. The species of this clade usually have a strong costa and a long seta, the seta often being mammillose in its upper part. The peristome types in this clade are mixed, since both reduced and unreduced types are found. Several neckeraceous genera that were recognised on a morphological basis are polyphyletic (e.g. Neckera, Homalia, Thamnobryum, Porotrichum). Ancestral state reconstructions revealed that currently used diagnostic traits, such as the leaf asymmetry and costa strength are highly homoplastic. Similarly, the reconstructions revealed that the 'reduced' sporophyte features have evolved independently in each of the three clades. Earlier phylogenetic studies, including species belonging to the Neckeraceae, have indicated that this pleurocarpous moss family shares a strongly supported sister group relationship with the Lembophyllaceae, but the family delimitation of the former needs adjustment. To test the monophyly of the Neckeraceae, as well as to redefine the family circumscription and to pinpoint its phylogenetic position in a larger context, a phylogenetic study based on molecular data was carried out. Sequence data were compiled, combining data from all three genomes: nuclear ITS1 and 2, plastid trnS-rps4-trnT-trnL-trnF and rpl16, and mitochondrial nad5 intron. The Neckeraceae have sometimes been divided into the two families, Neckeraceae and Thamnobryaceae, a division rejected here. Both parsimony and Bayesian analyses of molecular data revealed that the family concept of the Neckeraceae needs several further adjustments, such as the exclusion of some individual species and smaller genera as well as the inclusion of the Leptodontaceae. Within the family three well-supported clades (A, B and C) can be distinguished. Members of clade A are mainly non-Asiatic and nontropical. Most species have a weak costa and immersed capsules with reduced peristomes (mainly Neckera spp.) and the teeth at the leaf margins are usually unicellular. Clade B members are also mainly non-Asiatic. They are typically fairly robust, distinctly stipilate, having a single, at least relatively strong costa, long setae (capsules exserted), and the peristomes are well developed or only somewhat reduced. Members of clade C are essentially Asiatic and tropical. The species of this clade usually have a strong costa and a long seta, the seta often being mammillose in its upper part. The peristome types in this clade are mixed, since both reduced and unreduced types are found. Several neckeraceous genera that were recognised on a morphological basis are polyphyletic (e.g. Neckera, Homalia, Thamnobryum, Porotrichum). Ancestral state reconstructions revealed that currently used diagnostic traits, such as the leaf asymmetry and costa strength are highly homoplastic. Similarly, the reconstructions revealed that the 'reduced' sporophyte features have evolved independently in each of the three clades.

Phylogenetic analyses of the Hypnales usually show the same picture of poorly resolved trees with a large number of polyphyletic taxa and low support for the few reconstructed clades. One odd clade, however, consisting of three genera that are currently treated either within the Leskeaceae (Miyabea) or Neckeraceae (Homaliadelphus and Bissetia), was retrieved in a previously published phylogeny based on chloroplast rbcL. In order to elucidate the reliability of the observed Homaliadelphus - Miyabea - Bissetia - clade (HMB-clade) and to reveal its phylogenetic relationships a molecular study based on a representative set of hypnalean taxa was performed. Sequence data from all three genomes, namely the ITS1 and 2 (nuclear), the trnS-rps4-trnT-trnL-trnF cluster (plastid), the nad5 intron (mitochondrial), were analyzed. Although the phylogenetic reconstruction of the combined data set was not fully resolved regarding the backbone it clearly indicated the polyphyletic nature of various hypnalean families, such as the Leskeaceae, Hypnaceae, Hylocomiaceae, Neckeraceae, Leptodontaceae and Anomodontaceae with respect to the included taxa. In addition the results favor the inclusion of the Leptodontaceae and Thamnobryaceae in the Neckeraceae. The maximally supported HMB-clade consisting of the three genera Homaliadelphus (2-3 species), Miyabea (3 species) and Bissetia (1 species) is resolved sister to a so far unnamed clade comprising Taxiphyllum aomoriense, Glossadelphus ogatae and Leptopterigynandrum. The well-resolved and supported HMB-clade, here formally described as the Miyabeaceae, fam. nov. is additionally supported by morphological characters such as strongly incrassate, porose leaf cells, a relatively weak and diffuse costa and the presence of dwarf males. The latter are absent in the Neckeraceae and the Leskeaceae. It is essentially an East Asian family, with one species occurring in North America. Phylogenetic analyses of the Hypnales usually show the same picture of poorly resolved trees with a large number of polyphyletic taxa and low support for the few reconstructed clades. One odd clade, however, consisting of three genera that are currently treated either within the Leskeaceae (Miyabea) or Neckeraceae (Homaliadelphus and Bissetia), was retrieved in a previously published phylogeny based on chloroplast rbcL. In order to elucidate the reliability of the observed Homaliadelphus - Miyabea - Bissetia - clade (HMB-clade) and to reveal its phylogenetic relationships a molecular study based on a representative set of hypnalean taxa was performed. Sequence data from all three genomes, namely the ITS1 and 2 (nuclear), the trnS-rps4-trnT-trnL-trnF cluster (plastid), the nad5 intron (mitochondrial), were analyzed. Although the phylogenetic reconstruction of the combined data set was not fully resolved regarding the backbone it clearly indicated the polyphyletic nature of various hypnalean families, such as the Leskeaceae, Hypnaceae, Hylocomiaceae, Neckeraceae, Leptodontaceae and Anomodontaceae with respect to the included taxa. In addition the results favor the inclusion of the Leptodontaceae and Thamnobryaceae in the Neckeraceae. The maximally supported HMB-clade consisting of the three genera Homaliadelphus (2-3 species), Miyabea (3 species) and Bissetia (1 species) is resolved sister to a so far unnamed clade comprising Taxiphyllum aomoriense, Glossadelphus ogatae and Leptopterigynandrum. The well-resolved and supported HMB-clade, here formally described as the Miyabeaceae, fam. nov. is additionally supported by morphological characters such as strongly incrassate, porose leaf cells, a relatively weak and diffuse costa and the presence of dwarf males. The latter are absent in the Neckeraceae and the Leskeaceae. It is essentially an East Asian family, with one species occurring in North America. Phylogenetic analyses of the Hypnales usually show the same picture of poorly resolved trees with a large number of polyphyletic taxa and low support for the few reconstructed clades. One odd clade, however, consisting of three genera that are currently treated either within the Leskeaceae (Miyabea) or Neckeraceae (Homaliadelphus and Bissetia), was retrieved in a previously published phylogeny based on chloroplast rbcL. In order to elucidate the reliability of the observed Homaliadelphus - Miyabea - Bissetia - clade (HMB-clade) and to reveal its phylogenetic relationships a molecular study based on a representative set of hypnalean taxa was performed. Sequence data from all three genomes, namely the ITS1 and 2 (nuclear), the trnS-rps4-trnT-trnL-trnF cluster (plastid), the nad5 intron (mitochondrial), were analyzed. Although the phylogenetic reconstruction of the combined data set was not fully resolved regarding the backbone it clearly indicated the polyphyletic nature of various hypnalean families, such as the Leskeaceae, Hypnaceae, Hylocomiaceae, Neckeraceae, Leptodontaceae and Anomodontaceae with respect to the included taxa. In addition the results favor the inclusion of the Leptodontaceae and Thamnobryaceae in the Neckeraceae. The maximally supported HMB-clade consisting of the three genera Homaliadelphus (2-3 species), Miyabea (3 species) and Bissetia (1 species) is resolved sister to a so far unnamed clade comprising Taxiphyllum aomoriense, Glossadelphus ogatae and Leptopterigynandrum. The well-resolved and supported HMB-clade, here formally described as the Miyabeaceae, fam. nov. is additionally supported by morphological characters such as strongly incrassate, porose leaf cells, a relatively weak and diffuse costa and the presence of dwarf males. The latter are absent in the Neckeraceae and the Leskeaceae. It is essentially an East Asian family, with one species occurring in North America.

In order to delimit and understand the evolution of the Meteoriaceae, we provide phylogenetic analyses using the internal transcribed spacer 2 (ITS2) of nuclear ribosomal DNA in combination with two plastid markers, trnL-F and psbT-H. In contrast to the widely used trnL-F region, the psbT-H gene cluster, coding for proteins of photosystem II, has been rarely used to address systematic questions among the different land plant lineages. To overcome the problem of potential ambiguous alignments of non-coding DNA regions, the data were independently analyzed using direct optimization. The comparison and evaluation of the obtained results showed that the inferred cladograms based on the different phylogenetic approaches are very similar, with only minor differences. In combination with morphological characters, generic relationships as well as taxonomic and nomenclatural problems, especially regarding the key genera Meteorium and Papillaria are discussed in detail. New insights into generic relationships of the Meteoriaceae are provided, such as the exclusion of the monospecific southern South American genera Ancistrodes and Cryphaeophilum, which are subsequently transferred to the Hookeriaceae and Cryphaeaceae, respectively. Phylogenetic reconstructions using maximum likelihood as well as parsimony approaches reveal that at the familial level the Meteoriaceae s. l. are polyphyletic, if the formerly recognized "Trachypodaceae" are considered as a separate family. Based on our results we favor the synonym of the Trachypodaceae with the Meteoriaceae.

The taxonomic position of Rhynchostegiella acicula , a local endemic of Shaanxi Province of China, is investigated by means of cladistic analyses employing nrITS sequences. The analyses show that R. acicula does not belong to Rhynchostegiella s.str. (Helicodontioideae) but is resolved within the Homalothecioideae as sister to Eurhynchiastrum , from which it differs by a soft and slender habit; narrow lanceolate and acuminate leaves; a percurrent costa; and an autoicous condition. It differs from Brachytheciastrum and Brachythecium in a longly rostrate operculum, and from Homalothecium in a small plants that lack thick-walled basal laminal cells characteristic of this genus, as well as in an autoicous inflorescence. As a consequence, R. acicula is transferred into its own, monospecific genus Frahmiella Ignatov, Vanderpoorten & Wang You-fang, gen. nov.

The systematic position of the moss genus Acrocladium Mitten is reviewed and previous family placements are evaluated. A molecular dataset spanning five genomic regions from all three genomes was developed to address the relationships of Acrocladium . The results indicate a sister group relationship with Lepyrodontaceae, rather than with the Lembophyllaceae or Amblystegiaceae, as previously suggested. The strong differences in the morphologies of Lepyrodon and Acrocladium warrant recognition of the latter at family level: Acrocladiaceae R. Tangney, D. Quandt, S. Huttunen and M. Stech, fam. nov.

The presence of short inversions in non-coding plastid DNA is universal and an integral part of its evolution. We studied the effect of these structural changes in phylogenetic inference by measuring phylogenetic accuracy as a topological congruence among topologies inferred from plastid and nuclear sequences in ferns ( Lindsaea ) and mosses (Brachytheciaceae). We randomly replicated ten subsets of both groups and tested the performance of various coding schemes and phylogenetic methods. Our results demonstrate that short inversions do interfere with phylogenetic analysis, leading into incongruent topologies. The phylogenetic accuracy was increased by removing inversions or applying various coding schemes to them. Parsimony analyses resulted in greater accuracy than maximum likelihood or Bayesian analyses, but this was at least partly explained by slightly poorer resolution of parsimony trees. Due to the great importance of non-coding plastid DNA in species-level plant systematics and the frequency of structural rearrangements in it, we recommend these kinds of data be carefully explored before analysis, and the possible rearrangements to be optimized.