Tamás Pócs

World checklist of hornworts and liverworts

Abstract A working checklist of accepted taxa worldwide is vital in achieving the goal of developing an online flora of all known plants by 2020 as part of the Global Strategy for Plant Conservation. We here present the first-ever worldwide checklist for liverworts (Marchantiophyta) and hornworts (Anthocerotophyta) that includes 7486 species in 398 genera representing 92 families from the two phyla. The checklist has far reaching implications and applications, including providing a valuable tool for taxonomists and systematists, analyzing phytogeographic and diversity patterns, aiding in the assessment of floristic and taxonomic knowledge, and identifying geographical gaps in our understanding of the global liverwort and hornwort flora. The checklist is derived from a working data set centralizing nomenclature, taxonomy and geography on a global scale. Prior to this effort a lack of centralization has been a major impediment for the study and analysis of species richness, conservation and systematic research at both regional and global scales. The success of this checklist, initiated in 2008, has been underpinned by its community approach involving taxonomic specialists working towards a consensus on taxonomy, nomenclature and distribution.

Molecular insights into the phylogeny and subgeneric classification of Frullania Raddi (Frullaniaceae, Porellales)

With an estimated 300-375 species, Frullania is the largest genus of Porellales and forms a major clade of leafy liverworts. The cosmopolitan genus includes mostly epiphytes and represents an important component of the cryptogamic vegetation in various, especially tropical, habitats. There have been abundant changes and modifications to the infrageneric classification of Frullania, with up to fifteen subgenera and numerous sections solely based on morphology. Here we present the first molecular phylogeny of Frullania using four markers (rbcL, psbA, trnL-trnF region of cp DNA, nr5.8S-ITS-2 region) and 171 accessions from throughout the range. The molecular data provide evidence for the monophyly of several subgenera and support intercontinental ranges of these clades. Previous subgeneric assignment for a suite of taxa based on morphological evidence is not supported by the molecular data. Representatives of the genera Amphijubula, Neohattoria and Schusterella are nested in robust subclades of Frullania. Basal relationships within Frullania are largely unsupported. Based on the outcome of the phylogenetic analyses we present a revised supraspecific classification and provide evidence for the monophyly of some morphological species. Disjunct distributional patterns within Frullania cannot be explained by Gondwanan vicariance.

Tramps, narrow endemics and morphologically cryptic species in the epiphyllous liverwort Diplasiolejeunea

Diplasiolejeunea is a pantropical, epiphytic genus of leafy liverworts that occurs from the lowlands to more than 4000m altitude. Phylogenetic analyses of a molecular dataset consisting of three markers (nuclear ribosomal ITS region, plastidic trnL-F region and rbcL gene) and 122 accessions (plus two outgroups, Colura and Cololejeunea) indicate that the evolutionary diversity of Diplasiolejeunea is underestimated by current morphology-based classification. Four morphologically semi-cryptic species have been recovered. The molecular phylogenies support a deep split into a Neotropical and a Paleotropical clade, the latter structured into Australasian, Asian and Afromadacascan lineages. Presented results confirm the ranges of two pantropical species (D. cavifolia, D. rudolphiana), provide evidence for dispersal from the Neotropics into the Paleotropics, indicate speciation along altitudinal gradients and demonstrate extensive morphological homoplasy. We propose a revised supraspecific classification of Diplasiolejeunea into a predominantly Paleotropical subgenus Physolejeunea and predominantly Neotropical subgenera Austrolejeuneopsis and Diplasiolejeunea, the former containing mainly epiphytic species, the latter mainly epiphylls. Several clades are supported by combinations of morphological character states, and could be assigned to sections at some later point. This is the first comprehensive phylogeny of a largely epiphyllous genus of liverworts.

New national and regional bryophyte records, 18

Feddes Repertorium 87: 188. 1976 [Euosmolejeunea cedercreutzii H.Buch & Perss., Commentationes Biologicae 8 (7): 9. 1941]. MADEIRA: on bark of moribund Erica, north side of Pico Topeiro, to east of Encumeada, ca 1150 m a.s.l., 8 April 2007, leg. T.L. Blockeel no. 36/102 (LISU, duplicate in Hb. Blockeel). AZORES: Terceira: in damp crevice of lava field, Misterio Negro, 10 km N.W. of Angra do Heroismo, UTM 75–87-, ca 650 m a.s.l., 15 July 1994, leg. J.W. Bates & R. Gabriel, no. 3498 (Hb. Bates); Terceira: on Laurus bark in steep forest, Terra Brava, about 1 km N.E. of Algar do Carvao, UTM 82–87-, ca 640 m a.s.l, 21 July 1994, leg. J.W. Bates & R. Gabriel, no. 3650 (Hb. Bates); Terceira: epiphyte on Juniperus in upper caldeira forest, Juncal, about 11 km N. of Angra do Heroismo, 22 July 1994, leg. J.W. Bates no. 3682 (Hb. Bates).

Molecular phylogeny of the leafy liverwort Lejeunea (Porellales): evidence for a neotropical origin, uneven distribution of sexual systems and insufficient taxonomy.

Background Lejeunea is a largely epiphytic, subcosmopolitan liverwort genus with a complex taxonomic history. Species circumscriptions and their relationships are subject to controversy; biogeographic history and diversification through time are largely unknown. Methodology and Results We employed sequences of two chloroplast regions (trnL-trnF, rbcL) and the nuclear ribosomal ITS region of 332 accessions to explore the phylogeny of the Harpalejeunea-Lejeunea-Microlejeunea complex. Lejeunea forms a well-supported clade that splits into two main lineages corresponding to L. subg. Lejeunea and L. subg. Crossotolejeunea. Neotropical accessions dominate early diverging lineages of both main clades of Lejeunea. This pattern suggests an origin in the Neotropics followed by several colonizations from the Neotropics into the Paleotropics and vice versa. Most Afro-Madagascan clades are related to Asian clades. Several temperate Lejeunea radiations were detected. Eighty two of the 91 investigated Lejeunea species could be identified to species level. Of these 82 species, 54 were represented by multiple accessions (25 para- or polyphyletic, 29 monophyletic). Twenty nine of the 36 investigated species of L. subg. Lejeunea were monoicous and 7 dioicous. Within L. subg. Crossotolejeunea, 15 of the 46 investigated species were monoicous and 31 dioicous. Some dioicous as well as some monoicous species have disjunct ranges. Conclusions/Significance We present the first global phylogeny of Lejeunea and the first example of a Neotropical origin of a Pantropical liverwort genus. Furthermore, we provide evidence for the Neotropics as a cradle of Lejeunea lineages and detect post-colonization radiations in Asia, Australasia, Afro-Madagascar and Europe. Dioicy/monoicy shifts are likely non-randomly distributed. The presented phylogeny points to the need of integrative taxonomical studies to clarify many Lejeunea binomials. Most importantly, it provides a framework for future studies on the diversification of this lineage in space and time, especially in the context of sexual systems in Lejeuneaceae.

Evidence for Rampant Homoplasy in the Phylogeny of the Epiphyllous Liverwort Genus Cololejeunea (Lejeuneaceae)

Abstract Rampant homoplasy can be a major challenge in the classification of land plants that have limited morphological differences and/or ecological diversity, such as liverworts and mosses. Here we present the first comprehensive molecular phylogeny of the derived liverwort genus Cololejeunea and its close relatives. We used three markers (trnL—F, rbcL, and nrITS) and 116 accessions representing the geographic range as well as the morphological diversity of this predominantly epiphyllous genus. The molecular data support three major lineages: Colura, Myriocoleopsis, and Cololejeunea. Myriocoleopsis species were resolved in a clade with Cololejeunea minutissima. Aphanolejeunea and Chondriolejeunea were both resolved as putative monophyletic groups, and nested in the Cololejeunea clade. Cololejeunea angustiflora was found to be sister to the clade comprising the remaining Cololejeunea species. Currently accepted subgenera of Cololejeunea were paraphyletic or polyphyletic. Six out of 30 species with multiple accessions were not supported as natural taxa. The current classifications of Cololejeunea have been influenced by frequent homoplasy of morphological characters and do not accurately reflect species relationships. A new classification for this genus is outlined. Our data did not provide evidence for ancient vicariance events where major clades and continents are associated, instead some evidence was recovered for recent intercontinental species range expansion.

Crown Group Lejeuneaceae and Pleurocarpous Mosses in Early Eocene (Ypresian) Indian Amber

Cambay amber originates from the warmest period of the Eocene, which is also well known for the appearance of early angiosperm-dominated megathermal forests. The humid climate of these forests may have triggered the evolution of epiphytic lineages of bryophytes; however, early Eocene fossils of bryophytes are rare. Here, we present evidence for lejeuneoid liverworts and pleurocarpous mosses in Cambay amber. The preserved morphology of the moss fossil is inconclusive for a detailed taxonomic treatment. The liverwort fossil is, however, distinctive; its zig-zagged stems, suberect complicate-bilobed leaves, large leaf lobules, and small, deeply bifid underleaves suggest a member of Lejeuneaceae subtribe Lejeuneinae (Harpalejeunea, Lejeunea, Microlejeunea). We tested alternative classification possibilities by conducting divergence time estimates based on DNA sequence variation of Lejeuneinae using the age of the fossil for corresponding age constraints. Consideration of the fossil as a stem group member of Microlejeunea or Lejeunea resulted in an Eocene to Late Cretaceous age of the Lejeuneinae crown group. This reconstruction is in good accordance with published divergence time estimates generated without the newly presented fossil evidence. Balancing available evidence, we describe the liverwort fossil as the extinct species Microlejeunea nyiahae, representing the oldest crown group fossil of Lejeuneaceae.

Towards a monophyletic classification of Lejeuneaceae IV: reinstatement of Allorgella, transfer of Microlejeunea aphanella to Vitalianthus and refinements of the subtribal classification

The classification of Lejeuneeae, the most speciose tribe of the largely epiphytic leafy liverwort family Lejeuneaceae, is deemed notoriously difficult and hampered by extensive morphological homoplasy. We employed sequences of two chloroplast regions (trnL-trnF, rbcL) and the nuclear ribosomal ITS region of several critical taxa to explore their phylogenetic position. Based on our phylogenetic analyses and accompanying morphological investigations, we transfer Microlejeunea aphanella to Vitalianthus, reinstate the genus Allorgella, transfer Vitalianthus from Drepanolejeuneinae to Lepidolejeuneinae, identify Metalejeunea as another element of Lepidolejeuneinae, and place Allorgella and Haplolejeunea in Echinolejeuneinae. We map selected morphological characters on the reconstructed phylogenetic hypothesis to demonstrate the importance of subgynoecial innovations and ocelli for the circumscription of genera and subtribes of Lejeuneeae.

BIOGEOGRAPHY OF THE CUBAN BRYOPHYTE FLORA

Summary A distribution pattern analysis of 377 Cuban liverworts and 386 mosses shows a relatively high rate of endemism (11.4% and 12.2%, respectively), which is comparable to the rate of endemism in Cuban ferns but much lower than phanerogamic endemism (51.4%). In the Cuban bryoflora neotropical and Caribbean elements as well as Antillean subendemics prevail. The species/genus ratio (4:1 for the liverworts and 2:7 for the mosses), and the relatively high insularity index (3:1 for liverworts and 5:0 for mosses) are indicative of long-term isolation of the Cuban bryophyte flora and the large size of the island.

Phylogeny and Classification of Lejeuneaceae Subtribe Cheilolejeuneinae (Marchantiophyta) Based on Nuclear and Plastid Molecular Markers

Abstract Cheilolejeuneinae is an early diverging lineage of Lejeuneaceae tribe Lejeuneeae with a pantropical distribution. The current phylogeny and classification of this subtribe is based on morphological and limited-sampling molecular studies. Here we present a molecular phylogeny of Cheilolejeuneinae and related lineages based on maximum parsimony and maximum likelihood analyses, as well as Bayesian inference of two chloroplast regions (trnL-F, trnG) and the nuclear ribosomal ITS1-5.8S-ITS2 region, to test the monophyly of this subtribe, and to re-evaluate the infrageneric classification of Cheilolejeunea. The results confirm the monophyly of Cheilolejeuneinae, which consists of one single genus, Cheilolejeunea. Aureolejeunea and Omphalanthus are nested in Cheilolejeunea and, consequently, all species (11) and varieties (2) accepted in these two genera are transferred to Cheilolejeunea. Based on morphological and molecular evidence a new infrageneric classification of Cheilolejeunea into nine sections is proposed: sect. Anomalolejeunea, sect. Cheilolejeunea, sect. Cyrtolejeunea, sect. Euosmolejeunea, sect. Leucolejeunea, sect. Paroicae, sect. Omphalanthus, sect. Strepsilejeunea and sect. Xenolejeunea. Cheilolejeunea decurviloba is resolved in a sister relationship to Pycnolejeunea densistipula and C. gaoi constitues a separate lineage remote from Cheilolejeuneinae.