Raymond E. Stotler

PHYLOGENY AND CLASSIFICATION OF THE MARCHANTIOPHYTA

Input from molecular phylogenetics in the past five years has substantially altered concepts of systematic relationships among liverworts. While these studies have confirmed the monophyly of phylum Marchantiophyta, they have demonstrated that many previously recognised ranks within the hierarchy are unnatural and in need of modification. Changes in the ranks of suborder and above have been proposed by various workers, but modifications in the circumscription of genera and families are still required. A comprehensive, phylogenetic classification scheme that integrates morphological data with molecular hypotheses is presented. The scheme includes diagnoses and publication citations for all names above the rank of genus. All currently recognised genera are listed alphabetically in their respective families; subfamilies are not indicated. Major modifications and novel alignments of taxa are thoroughly discussed, with pertinent references provided. Jungermanniaceae is redefined and Solenostomataceae fam. nov. is formally described to accommodate some of the genera excluded from it.

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.

A Revised Classification of the Anthocerotophyta and a Checklist of the Hornworts of North America, North of Mexico

Abstract A hornwort classification that recognizes 11 genera, comprising a hierarchy of two classes, three orders, and four families is proposed. The updated checklist of these bryophytes for North America has been expanded from 11 species in three genera to 17 species in five genera. Application of the genus name Anthoceros in the traditional sense is clarified.

Taxonomy and Classification

This chapter provides a description of all plant families and genera that include putative fully mycoheterotrophic species, excluding initial and partial mycoheterotrophs. The overview covers a total of 17 families, 101 genera, and ca. 880 species. For each family and genus (except for Orchidaceae) a short morphological description is provided followed by notes on taxonomy, distribution, evolution, and ecology. For most genera a line drawing of a representative species is provided. Included families are: Aneuraceae, Burmanniaceae, Corsiaceae, Ericaceae, Gentianaceae, Gleicheniaceae, Iridaceae, Lycopodiaceae, Ophioglossaceae, Orchidaceae, Petrosaviaceae, Podocarpaceae, Polygalaceae, Psilotaceae, Schizaeaceae, Thismiaceae, and Triuridaceae.

CONTRIBUTIONS TOWARD A BRYOFLORA OF CALIFORNIA III. KEYS AND ANNOTATED SPECIES CATALOGUE FOR LIVERWORTS AND HORNWORTS

ABSTRACT Keys are provided to 54 genera, 142 species, 6 infraspecific taxa, and 3 undescribed species of California liverworts and hornworts within 33 families. In addition, 3 genera and 16 species previously reported have been excluded from the California bryoflora. The keys emphasize gametophytic characters. Annotated comments for each species focus on what to look for and how to separate it from species with similar morphology. The worldwide and California distribution of each species is provided.

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.

A Molecular Study of the Simple Thalloid Liverwort Jensenia (Marchantiophyta, Pallaviciniaceae)

Abstract Five chloroplast coding and non-coding regions, one mitochondrial gene and its associated intron, and part of one nuclear ribosomal gene were sequenced, giving a total included aligned length of 7,845 bp. Samples from six collections, representing five species of Jensenia from South America, Malaysia, and New Zealand, four collections belonging to three species of Pallavicinia and one collection of Podomitrium phyllanthus comprised the ingroup, which was rooted on Symphyogyna marginata. Phylogenetic analysis under the maximum parsimony and maximum likelihood criteria support Jensenia as monophyletic. Within Jensenia, two monophyletic clades are resolved, one comprising the four Latin American collections and the other, the Malayasian and New Zealand species; however, this is based on only 16 parsimony informative characters. The lack of informative characters within Jensenia suggests that the divergence among taxa is relatively recent, rather than Gondwanan, as has been hypothesized based on geographical distribution. A corollary of this is that the present day distribution of Jensenia is likely the product of dispersal rather than vicariance.

Typifications in the Genus Fossombronia (Marchantiophyta)

Abstract Lectotypes are explicitly designated for 43 taxa in the genus Fossombronia Raddi, with neotypes designated for F. mauritanica Trab. and F. pusilla var. β capitata Nees. One name, F. pumila Dumort, is regarded as an orthographic error not based upon a specimen and thus, cannot be typified. The entries are listed alphabetically, each with the correct author citation, place of publication, and the herbarium of deposit. In addition, supporting comments are given for most selections. Fossombronia caespitiformis De Not. ex Rabenh. var. multispira Schiffn. is elevated to the rank of subspecies. Images of all type specimen labels may be viewed at: http://bryophytes.plant.siu.edu/fossombronia.html

Contributions toward a Monograph of Petalophyllum (Marchantiophyta)

The simple thalloid liverwort genus Petalophyllum J. G. C. Lehmann is removed from the Fossombroniaceae and sequestered in the new family Petalophyllaceae along with Sewardiella S. R. Kashyap, a monotypic genus endemic to the In- dian subcontinent. Unlike the Fossombroniaceae, the thallus in the Petalophyllaceae is never dis- sected, the archegonia are clustered rather than scattered, a pseudoperianth is formed rather than a caulocalyx, and the capsule wall is 3-4-layered with outer wall thickenings, which are lacking in the Fossombroniaceae. The North American rep- resentative of Petalophyllum previously considered conspecific with the European taxon P. ralfsii (W. M. Wilson) J. G. C. Lehmann is here described as a distinct species, P. americanum. This is based upon clear differences of the thallus, pseudoper- ianth, and the elater spirals. Furthermore, the ap- plication of the name P. australis Colenso to New Zealand plants is shown to be incorrect; those plants previously referred there represent a new species, P. hodgsonii. This species resembles P. ralfsii, but differs from it in the broad, strongly flat- tened thallus midrib and structure of the female inflorescence.

On the taxonomic status of the enigmatic Phycolepidoziaceae (Marchantiophyta: Jungermanniales) with description of a new species, Phycolepidozia indica

The monospecific Phycolepidoziaceae with the single neotropical species Phycolepidozia exigua is a highly specialized leafy liverwort without vegetative leaves. The extreme reduction of morphological and anatomical characters of Phycolepidozia has caused uncertainties as to the systematic position of the genus and family. In 2012, a second species of Phycolepidozia was detected in the Western Ghats, South India. The Indian plant differs from P. exigua in several respects and is described here as P. (subg. Metaphycolepidozia) indica Gradst., J.-P.Frahm & U.Schwarz. Differences include the massive stem of P. indica, the larger perianth with a crenate, 3-lobed mouth, and the epidermis of the capsule wall made up of non-tiered cells with nodular thickenings on both longitudinal and transverse walls. A phylogenetic analysis using four different chloroplast regions (psbA, psbT, rps4, rbcL) of P. indica and putatively related groups shows that Phycolepidozia is nested within the leafy liverwort family Cephaloziellaceae. Consequently, Phycolepidoziaceae is placed in the synonymy of Cephaloziellaceae. The discovery of P. indica adds a further example to the list of amphi-Pacific tropical disjunctions in bryophytes.