John J. Engel

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.

Goodbye or welcome Gondwana? – insights into the phylogenetic biogeography of the leafy liverwort Plagiochila with a description of Proskauera, gen. nov. (Plagiochilaceae, Jungermanniales)

Molecular phylogenies based on chloroplast gene rps4 sequences and nuclear ribosomal ITS sequences have been generated to investigate relationships among species and putative segregates in Plagiochila (Plagiochilaceae), the largest genus of leafy liverworts. About a fourth of the ca. 450 accepted binomials of Plagiochilaceae are included in these phylogenetic analyses, several represented by multiple accessions. A clade with Chiastocaulon, Pedinophyllum, and Plagiochilion is placed sister to a clade with numerous accessions of Plagiochila. Plagiochila pleurata and P. fruticella are resolved sister to the remainder of Plagiochilaceae and transferred to the new Australasian genus Proskauera which differs from all other Plagiochilaceae by the occurrence of spherical leaf papillae. The historical biogeography of Plagiochilaceae is explored based on the reconstructions of the phylogeny, biogeographic patterns and diversification time estimates. The results indicate that the current distribution of Plagiochilaceae cannot be explained exclusively by Gondwanan vicariance. A more feasible explanation of the range is a combination of short distance dispersal, rare long distance dispersal events, extinction, recolonization and diversification.

Extant diversity of bryophytes emerged from successive post-Mesozoic diversification bursts

Unraveling the macroevolutionary history of bryophytes, which arose soon after the origin of land plants but exhibit substantially lower species richness than the more recently derived angiosperms, has been challenged by the scarce fossil record. Here we demonstrate that overall estimates of net species diversification are approximately half those reported in ferns and ∼30% those described for angiosperms. Nevertheless, statistical rate analyses on time-calibrated large-scale phylogenies reveal that mosses and liverworts underwent bursts of diversification since the mid-Mesozoic. The diversification rates further increase in specific lineages towards the Cenozoic to reach, in the most recently derived lineages, values that are comparable to those reported in angiosperms. This suggests that low diversification rates do not fully account for current patterns of bryophyte species richness, and we hypothesize that, as in gymnosperms, the low extant bryophyte species richness also results from massive extinctions.

Austral Hepaticae. 35. A Taxonomic and Phylogenetic Study of Telaranea (Lepidoziaceae), with a Monograph of the Genus in Temperate Australasia and Commentary on Extra-Australasian Taxa

Abstract Telaranea is the third largest genus of Lepidoziaceae, after Bazzania (over 450 species) and Lepidozia (over 300 species). In this monograph, 98 species of Telaranea are recognized, 62 of which are extra-Australasian. Included are 11 new species, 2 new varieties, 27 new combinations (25 species and one variety), and one new name. New taxa are fully described and illustrated. Two subgenera of Telaranea are recognized (subg. Acrolepidozia, subg. Telaranea), and 7 sections within subg. Telaranea (sect. Neolepidozia (Fulf. & J. Tayl.) Engel & Merr. comb. nov., sect. Cancellatae Engel & Merr., sect. Ceraceae Engel & Merr. sect. nov., sect. Tricholepidozia (Schust.) Engel & Merr. comb. nov., sect. Transversae Engel & Merr., sect. Telaranea, sect. Tenuifoliae (Schust.) Engel & Merr. comb. nov.). A major part of this work is devoted to a phylogenetic study of the genus Telaranea, with a discussion of character evolution in the genus and a reconstruction of ancestral character states. An ancestral area analysis was also undertaken, using phylogeny to trace the geographical history of members of the genus. Differing concepts of phylogeny as applied to leafy hepatics are also briefly considered. The phylogenetic analysis included 56 taxa and 32 characters, with representative species of Lepidozia, Kurzia, and Arachniopsis as out-groups. Cladograms obtained from parsimony analyses of the full data set and subsets of taxa, as well as distance (NJ) trees, were strikingly similar in topology. Monophyly of Telaranea is supported by the possession of a hyaloderm (enlarged, usually hyaline stem cortical cells) and an undivided first branch underleaf. Progressive reduction in morphological complexity as a central evolutionary tendency in the genus is strongly supported. The phylogeny indicates that Kurzia, as defined by the presence of Microlepidozia-type branching, is polyphyletic, and 6 species previously assigned to Kurzia are transferred to Telaranea. No support was found for maintaining Arachniopsis as a distinct genus, and 6 species of the genus are transferred to Telaranea. The new genus Amazoopsis Engel & Merr. is described, with 3 species: A. diplopoda (Pócs) Engel & Merr. comb. nov., A. dissotricha (Spruce) Engel & Merr. comb. nov., and A. gracilis Engel & Merr. sp. nov. The status of Monodactylopsis (Schust.) Schust. is discussed; the genus includes M. monodactyla (Spruce) Schust. and M. minima Schust. ex Engel & Merr., sp. nov. Paracromastigum vastilobum (Steph.) Engel & Merr. is a new combination. Keys to the subgenera and sections of Telaranea, the Australasian species, as well as the southern South American species of Telaranea, and the species of Telaranea sect. Tenuifoliae are included.

Studies of New Zealand Hepaticae. 1–6

Six New Zealand species are treated. 1. The correct basionym reference forBazzania hochstetteri (Reichardt) Hodgson is Reichardt (1866). A key to the New Zealand vittate species ofBazzania is included. 2. The correct citation forBazzania mittenii (Stephani) Stephani is given. 3.Lepidozia leptodictyon Herzog is placed in the synonymy ofPseudocephalozia lepidozioides R. Schuster. 4.Metahygrobiella chilensis Engel & Schuster is a new name forCephalozia heteroica Schuster & Engel. 5.Metzgeria alpina Schuster & Engel is described as new. 6. The correct citation forPachyglossa Herzog is Herzog (1952).

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.

The systematic identity of Chiloscyphus trichocoleoides, a new liverwort species from New Zealand, uncovered by morphological and molecular evidence

Abstract A new liverwort species from Western Nelson Ecological Province of New Zealand known from a single sterile specimen is described. It is unusual in having well-developed stem paraphyllia, and the leaves and underleaves are divided with three levels of ciliate division in a pinnate manner. This morphology suggested membership of Trichocoleaceae or Trichotemnomaceae, but other features of the plant made placement in either family seem unnatural. We determined its position by conducting phylogenetic analyses of rbcL, rps4 and trnL–F sequence datasets from 35 exemplars, and further confirmation of the position was carried out by an extended analysis using rbcL sequence dataset of 59 exemplars. The surprising result is that it belongs within the genus Chiloscyphus, where it is unique in these features. We describe this new species as Chiloscyphus trichocoleoides. Placing the new species in Chiloscyphus has broadened the generic concept of Chiloscyphus, and its phylogenetic relationship within and to other genera such as Clasmatocolea and Heteroscyphus needs further investigation. The results also show that the same morphological features seen in Chiloscyphus and in Trichocoleaceae and Trichotemnomaceae have developed independently.

Studies on Geocalycaceae XIV. Physotheca J. J. Engel & Gradst., a new genus of Hepaticae from Ecuador, belonging to a new subfamily, Geocalycaceae subfam. Physothecoideae J. J. Engel & Gradst.

Physotheca autoica J. J. Engel & Gradst. is a liverwort genus and species new to science and is endemic to the Andes of southern Ecuador. The new genus, while having several characters also found in the family Geocalycaceae, differs from members of that family in a number of important respects, which are discussed. The mode of growth of Physotheca is unusual, and occurs strictly by repeated innovations originating from below the bracts or bracteoles of the gynoecium. The inflated, bladder-like perianths are noteworthy, and have a shape and form that is remarkably similar to the sterile perianths of the unrelated genus Pleurozia. Capsule wall anatomy is also unusual, the walls are 7-8-stratose, and the outer layer, which has both one-phase and two-phase development represented, readily collapses. The large spores (to nearly 39 gm) are echinate and nearly three times the elater width, and the elaters, which frequently uncoil, adhere in considerable number to the inner face of the capsule wall. The best placement of Physotheca is within the family Geocalycaceae. Due to the unusual combination of these and other characters, the genus does not fit into any of the known subfamilies of Geocalycaceae and must be placed in a new subfamily, Physothecoideae J.J. Engel & Gradst.

The resurrection of Neohattoria Kamim. (Jubulaceae, Marchantiophyta): a six decade systematic conflict resolved through a molecular perspective

Abstract The systematic placement of Frullania herzogii has been contentious since its description six decades ago. Over the years it has been interpreted as either a member of the genus Frullania or segregated into its own genus, Neohattoria, due to morphological similarities with both Frullania and Jubula. Here we provide molecular evidence that supports the recognition of the genus Neohattoria and its inclusion within the Jubulaceae, together with Jubula and Nipponolejeunea. Jubulaceae are placed sister to Lejeuneaceae rather than to the monogeneric Frullaniaceae.

The status and phylogeography of the liverwort genus Apometzgeria Kuwah. (Metzgeriaceae)

Abstract Three species have been formerly segregated from Metzgeria in the genus Apometzgeria. One of the species, A. frontipilis, is endemic to South America but the second, A. pubescens, is currently understood as having a bipolar range with populations across the Holarctic and in southern South America. The third species, A. longifrondis, was described from China (and is not included in this study). Species of bryophytes that range across continents and have little or no morphological variation among populations may nevertheless harbor morphologically cryptic genetic lineages. We used nuclear and plastid sequence data to examine the phylogenetic relationship between Apometzgeria and Metzgeria, and phylogeographic patterns in taxa assigned to Apometzgeria. Two species often assigned to Apometzgeria are phylogenetically embedded within Metzgeria in two separate clades, one comprising all Holarctic A. pubescens and a second with A. pubescens from South America and all accessions of A. frontipilis. Phylogenetic and haplotype analyses reveal a lack of phylogeographic structure among A. pubescens plants from throughout its Holarctic distribution. However, A. pubescens in South America is more closely related to A. frontipilis and species of Metzgeria from South America than to any A. pubescens from the Northern Hemisphere. Thus, A. pubescens is Holarctic in distribution and morphologically similar plants form a divergent lineage in South America. Our results do not support Apometzgeria as a separate genus in the Metzgeriaceae.