Matt Von Konrat

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.

Morphologically cryptic biological species within the liverwort Frullania asagrayana

UNLABELLED PREMISE OF THE STUDY The Frullania tamarisci complex includes eight Holarctic liverwort species. One of these, F. asagrayana, is distributed broadly throughout eastern North America from Canada to the Gulf Coast. Preliminary genetic data suggested that the species includes two groups of populations. This study was designed to test whether the two groups are reproductively isolated biological species. • METHODS Eighty-eight samples from across the range of F. asagrayana, plus 73 samples from one population, were genotyped for 13 microsatellite loci. Sequences for two plastid loci and nrITS were obtained from 13 accessions. Genetic data were analyzed using coalescent models and Bayesian inference. • KEY RESULTS Frullania asagrayana is sequence-invariant at the two plastid loci and ITS2, but two clear groups were resolved by microsatellites. The two groups are largely reproductively isolated, but there is a low level of gene flow from the southern to the northern group. No gene flow was detected in the other direction. A local population was heterogeneous but displayed strong genetic structure. • CONCLUSIONS The genetic structure of F. asagrayana in eastern North America reflects morphologically cryptic differentiation between reproductively isolated groups of populations, near-panmixis within groups, and clonal propagation at local scales. Reproductive isolation between groups that are invariant at the level of nucleotide sequences shows that caution must be exercised in making taxonomic and evolutionary inferences from reciprocal monophyly (or lack thereof) between putative species.

Volatile components of selected species of the liverwort genera Frullania and Schusterella (Frullaniaceae) from New Zealand, Australia and South America: a chemosystematic approach

The volatile components of 25 taxa of the liverwort family Frullaniaceae from New Zealand, Australia and South America have been analyzed by GC-MS. The present Frullania species are chemically divided into five major types: (1) sesquiterpene lactones, (2) sesquiterpene lactones-bibenzyls, (3) bibenzyls, (4) 2-alkanone and (5) triterpene types; the latter two chemo-types are newly proposed for the genus. Schusterella chevalierii, belonging to the Frullaniaceae, is closely related chemically to the sesquiterpene lactone type of the Frullania species since it elaborates two eudesmanolides, beta-cyclocostunolide and dihydro-beta-cyclocostunolide as major components.

Integrative taxonomy resolves the cryptic and pseudo-cryptic Radula buccinifera complex (Porellales, Jungermanniopsida), including two reinstated and five new species

Abstract Molecular data from three chloroplast markers resolve individuals attributable to Radula buccinifera in six lineages belonging to two subgenera, indicating the species is polyphyletic as currently circumscribed. All lineages are morphologically diagnosable, but one pair exhibits such morphological overlap that they can be considered cryptic. Molecular and morphological data justify the re-instatement of a broadly circumscribed ecologically variable R. strangulata, of R. mittenii, and the description of five new species. Two species Radula mittenii Steph. and R. notabilis sp. nov. are endemic to the Wet Tropics Bioregion of north-east Queensland, suggesting high diversity and high endemism might characterise the bryoflora of this relatively isolated wet-tropical region. Radula demissa sp. nov. is endemic to southern temperate Australasia, and like R. strangulata occurs on both sides of the Tasman Sea. Radula imposita sp. nov. is a twig and leaf epiphyte found in association with waterways in New South Wales and Queensland. Another species, R. pugioniformis sp. nov., has been confused with Radula buccinifera but was not included in the molecular phylogeny. Morphological data suggest it may belong to subg. Odontoradula. Radula buccinifera is endemic to Australia including Western Australia and Tasmania, and to date is known from south of the Clarence River on the north coast of New South Wales. Nested within R. buccinifera is a morphologically distinct plant from Norfolk Island described as R. anisotoma sp. nov. Radula australiana is resolved as monophyletic, sister to a species occurring in east coast Australian rainforests, and nesting among the R. buccinifera lineages with strong support. The molecular phylogeny suggests several long-distance dispersal events may have occurred. These include two east-west dispersal events from New Zealand to Tasmania and south-east Australia in R. strangulata, one east-west dispersal event from Tasmania to Western Australia in R. buccinifera, and at least one west-east dispersal from Australia to New Zealand in R. australiana. Another west-east dispersal event from Australia to Norfolk Island may have led to the budding speciation of R. anisotoma. In contrast, Radula demissa is phylogeographically subdivided into strongly supported clades either side of the Tasman Sea, suggesting long distance dispersal is infrequent in this species.

Phylogenetic Relationships and Morphological Evolution in a Major Clade of Leafy Liverworts (Phylum Marchantiophyta, Order Jungermanniales): Suborder Jungermanniineae

Abstract— The suborder Jungermanniineae of the Jungermanniales is a major lineage of leafy liverworts, recognized in recent classifications to include 15 families. Gametophytes within the suborder are morphologically diverse, but commonly anisophyllous to distichous, usually with succubous, rarely transverse or incubuous, leaf insertions. Sporophytes are frequently, but not universally, enclosed by stemderived perigynia or coelocaules, often accompanied by perianth reduction or loss and some level of geocauly or marsupial development. We herein provide the first comprehensive molecular phylogeny of this geographically widespread suborder, using sequences generated from one nuclear (rpb2), two mitochondrial (nad1 and rps3), and seven plastid (atpB, psbA, psbT-H, rbcL, rps4, trnG and trnL) loci, sampled from 279 accessions representing 163 species in 57 genera. Ancestral states were reconstructed for 14 morphological characters generally considered taxonomically diagnostic for families in the suborder. Our phylogenetic analyses support the return of Leiomylia (=Mylia anomala) to the Myliaceae, removal of Myliaceae from the Jungermanniineae, and validation of the monogeneric suborder Myliineae subord. nov. to house it. Eighteen families are recognized within the Jungermanniineae, nine of which are monogeneric; namely, Endogemmataceae, Harpanthaceae, Gyrothyraceae, Arnelliaceae, Saccogynaceae, Geocalycaceae, Jackiellaceae, Notoscyphaceae stat. nov., and Trichotemnomaceae. The generic compositions of other families are modified as follows: Saccogynidium is transferred from Geocalycaceae to a newly named subfamily of Acrobolbaceae, Acrobolbaceae subf. Saccogynidioideae, and one other subfamily of the Acrobolbaceae is validated, Acrobolbaceae subf. Austrolophozioideae; Hygrobiella is included in Antheliaceae (previously in Cephaloziaceae or its own family); Jungermanniaceae is broadened to include Mesoptychiaceae and Delavayellaceae; Cryptocoleopsis and Nardia are transferred from Solenostomataceae to Gymnomitriaceae; Gottschelia, Herzogobryum, and Nothogymnomitrion are excluded from the Jungermanniineae; Solenostomataceae is recognized to include Solenostoma, Arctoscyphus, Cryptocolea, and Diplocolea. Additional nomenclatural changes include recognizing Horikawaella as a synonym of Solenostoma and Apomarsupella as a synonym of Gymnomitrion, establishing two new subgenera of Solenostoma, Solenostoma subg. Metasolenostoma and Solenostoma subg. Eucalyx, and transferring Jungermannia conchata to Cephalozia. Morphological character state reconstructions identify dioecious inflorescences, gametangia on leading stems, flagelliform or stoloniferous branches absent, dorsal leaf insertions not overlapping the stem midline, large underleaves, and lack of gemmae as ancestral within the Jungermanniineae. All morphological characters appear to be moderately to highly homoplasious within the suborder.

Frullania knightbridgei, a new liverwort (Frullaniaceae, Marchantiophyta) species from the deep south of Aotearoa-New Zealand based on an integrated evidence-based approach

Abstract Frullania is a large and taxonomically complex genus. A new liverwort species, Frullania knightbridgei sp. nov. from southern New Zealand, is described and illustrated. The new species, and its placement in Frullania subg. Microfrullania, is based on an integrated evidence-based approach derived from morphology, ecology, experimental growth studies of plasticity, as well as sequence data. Diagnostic characters associated with the leaf and lobule cell-wall anatomy, oil bodies, and spore ultra-structure distinguish it from all other New Zealand species of Frullania. A critical comparison is also made between Frullania knightbridgei and morphologically allied species of botanical regions outside the New Zealand region and an artificial key is provided. The new species is similar to some forms of the widespread Australasian species, Frullania rostrata, but has unique characters associated with the lobule and oil bodies. Frullania knightbridgei is remarkably interesting in comparison with the majority of Frullania species, and indeed liverworts in general, in that it is at least partially halotolerant. Maximum parsimony and maximum likelihood analyses of nuclear ribosomal ITS2 and plastidic trnL-trnF sequences from purported related speciesconfirms its independent taxonomic status and corroborates its placement within Frullania subg. Microfrullania.

Frullania chevalieri (Jubulaceae) in New Zealand, with a reassessment of Schusterella

Abstract Frullania chevalieri (R. M. Schust.) R. M. Schust., previously only known from New Caledonia, is formally reported for the first time for New Zealand. This species represents an important element in the Jubulaceae because it is morphologically allied to a small group of species historically assigned to Schusterella S. Hatt. et al., which has a complicated and controversial taxonomic history. There has been no critical reassessment of Schusterella for over 30 years since the genus was first proposed and this has led to competing contemporary classifications systems. However, chemical and morphological evidence indicates very strongly that Frullania chevalieri and its allies should indeed be retained in the genus Frullania. Frullania chevalieri is described and illustrated in detail and a brief synopsis of morphologically allied species is made; characters associated with the sporophyte and oil-bodies are reported for the first time. The distinguishing features of F. chevalieri and its close morphological allies include the small plant size coupled with the dentate margins of the leaf-lobe.

Land of wood and water: A checklist of liverworts and hornworts of Jamaica

Abstract The first ever liverwort and hornwort checklist is provided for Jamaica. We report 332 taxa in 314 species, 14 of which are invalid, illegitimate or in orphaned genera but we do not know where to place them. An additional 16 of the reported taxa are rejected as erroneously reported from Jamaica. The list is based on over 270 literature references, including monographs, regional studies, and molecular investigations. Except for a few early works in the 18th and early 19th century, and a scattering of other studies, the liverwort and hornwort flora of Jamaica has received negligible attention, despite its apparent diversity. The knowledge of the liverwort and hornwort flora, both in Jamaica and regionally in the Caribbean, remains so poor it is too early to reliably comment about range-restricted or endemic species. We predict that a number of new records remain to be reported, based on regionally widespread species that have been recorded elsewhere. On the other hand, it is envisaged that a number of the taxa presented here maybe synonyms of other taxa as they are taxonomically insufficiently known. As a consequence, Anastrophyllum harrisanum Steph. is reduced to a new synonym of A. tubulosum (Nees) Grolle.

Chapter Nine: Early Land Plants Today: Liverwort Species Diversity and the Relationship with Higher Taxonomy and Higher Plants

Abstract Liverworts (Marchantiophyta) form a conspicuous and important component in many terrestrial ecosystems throughout the world. Despite of their significance and abundance, studies of various aspects on global plant species richness and distribution patterns have almost exclusively focused on vascular plants. Yet, comprehensive studies of liverwort plant diversity have many implications and broad applications. We use a preliminary dataset that unites liverwort nomenclature, taxonomy, and geography based on some 60,000 records. Global maps are produced for the taxonomic ranks of species, genus, and family to provide a rapid guide of diversity across political units worldwide. The usefulness of higher level taxon analyses was investigated at the genus and family level to test the relationship with species richness. A reduced set of taxonomic ranks other than species has been proved to be useful for rapid and cost-effective assessment of biodiversity. We provide the first examination of how well this method performs for liverworts. Generic richness was slightly more accurately related to species richness than that of families, indicating surrogacy at this taxonomic level as a promising approach for the prediction of liverwort species richness. Finally, given the fact that vascular plant diversity and distribution patterns are often given high consideration in evaluating global networks of protected areas and biodiversity hotspots, we present a comparison of centers of species richness between liverworts and vascular plants. Several regions of high liverwort species richness lie outside the highest centers of vascular plant species richness. We conclude with ideas for future studies of the dataset, which has many exciting implications and applications for the study of liverwort distribution and diversity patterns.

New records, replacements, reinstatements and four new species in the Radula parvitexta and R. ventricosa species groups (Jungermanniopsida) in Australia: cases of mistaken identity

Abstract. Various published hypotheses regarding circumscription and relationships of species within the Radula parvitexta and R. ventricosa species groups were tested using molecular data from three chloroplast markers. The phylogeny resolves five clades within the R. parvitexta species group in Australia, which proves polyphyletic across two subgenera, or three subgenera if R. madagascariensis is included. One clade represents an undescribed species, R. psychosis sp. nov., one corresponds to R. madagascariensis, a new record for Australia, the others to R. ratkowskiana, R. tasmanica and R. robinsonii. R. ratkowskiana is reinstated from synonymy of R. tasmanica, and R. parvitexta is placed into synonymy of R. robinsonii. A second new species belonging to the R. parvitexta species group, R. kilgourii sp. nov., is described; however, it was not included in the phylogeny. Three clades were resolved within the R. ventricosa species group in Australia, which is nested within subg. Metaradula. These clades corresponded to R. jovetiana, R. loriana, which is reinstated from synonymy of R. ventricosa, and two new species, namely, R. myriopoda sp. nov. and R. forficata sp. nov. R. ventricosa is excluded from the Australian flora, because all material is referrable to R. loriana. R. forficata and R. kilgourii had not been collected before the present study. R. myriopoda and R. jovetiana exhibit overlap in morphology of the sterile gametophyte and can be reliably separated only on characters associated with the perianth mouth. They can be considered semicryptic species, and would not have been recognised independent of fieldwork and molecular investigations conducted as part of the present study.