Hiroyuki Akiyama

Ordinal Phylogeny within the Hypnobryalean Pleurocarpous Mosses Inferred from Cladistic Analyses of Three Chloroplast DNA Sequence Data Sets: trnL-F, rps4, and rbcL

Abstract Classification of families of hypnobryalean mosses into the Hypnales, Leucodontales, and Hookeriales has been taxonomically difficult. Several researchers have sequenced different genes for independent phylogenetic studies of these three pleurocarp groups. Our goal is to summarize available molecular data and compile the largest data set to infer phylogenetic relationships among families as basis for classification at ordinal level. Sequences of rbcL, trnL-F, and rps4 loci for 38 exemplars of most families of Hypnales, Leucodontales, and Hookeriales were analyzed to evaluate whether or not each of the three orders is monophyletic. Cladistic analyses of combined sequences, using five taxa in the Bryales as outgroups, reveal a robust clade (decay > 5) including all hypnobryalean pleurocarps. Within this group, one clade (decay = 2) includes only taxa of the Hookeriales, and is sister to a large monophyletic group (Hypnales sensu lato) containing all other taxa (decay = 2) previously in the Leucodontales and Hypnales. These relationships suggest that the ordinal level taxonomy needs to be reconsidered since major lineages detected do not correspond to the traditional Leucodontales or Hypnales. These two orders are not supported by any molecular evidence from rbcL, trnL-F, or rps4, either analyzed singly or in different combinations. Additionally, present results indicate the need for changes to the current system of three suborders of Hypnales and four of the Leucodontales. Phylogenetic reconstructions based on molecular data emphasize the need for a re-examination of the taxonomic relevance of morphological characters and corroborate previous interpretations of sporophytic morphological similarities as multiple transitions to similar solutions to epiphytism among the pleurocarps.

Molecular Phylogeny of the Suborder Leucodontineae (Musci; Leucodontales) Inferred from rbcL Sequence Data

rbcL sequence data (1324 bp on average). Our analyses indicate (1) that Antitrichia is distantly related to the other members of Leucodontaceae and should be excluded from the family, (2) that Cryphaeaceae form a monophyletic clade, not with Anomodontaceae and Leptodontaceae, but with Leucodontaceae, refuting the placement of Leucodontaceae and Cryphaeaceae in different superfamilies, (3) that Forsstroemia, variously classified in Leucodontaceae, Cryphaeaceae or Leptodontaceae, forms a monophyletic clade with Neckera (Neckeraceae), and (4) that the presumed monophyly of Anomodon and that of Anomodontaceae are not supported.

Morphology and Taxonomic Significance of Dormant Branch Primordia, Dormant Buds, and Vegetative Reproductive Organs in the Suborders Leucodontineae and Neckerineae (Musci, Isobryales)

The manner of branch development and shape of vegetative reproductive organs were examined for 19families, 101 genera, and 227 species in the suborders Leucodontineae and Neck- erineae (Musci). Two types were recognized in the manner of branch development. They are distin- guished from each other by the nature of dormant condition: One with swollen primordia (type A) and the other with buds (type S). Filamentous pseudoparaphyllia always accompanied type A primordia, whilefoliose pseudoparaphyllia were always with type S. The presence ofscale-like leaves overarching buds was a good marker for recognizing the type S with ease. Each family and genus, with few exceptions, proved to be stable in the manner ofbranch development and shape of vegetative reproductive organs. Therefore, the two-character combination can be used as a new clue for the taxonomy of higher categories. Five genera, which are not closely related, have paraphyllia. The presence ofparaphyllia is correlated with the type S of branch development, and has no taxonomic significance. The familial affinities of two genera are considered; Henicodium is a monotypic genus and classified in the Pterobryaceae, and Penzigiella in the Neckeraceae. Several families of the Leucodontineae have long

Clastobryopsis imbricata (Pylaisiadelphaceae) sp. nov. from Doi Inthanon, northern Thailand

Abstract Clastobryopsis imbricata sp. nov. (Pylaisiadelphaceae, Musci) is described on the basis of specimens collected at middle elevations of Doi (Mt.) Inthanon, northern Thailand. The species is characterized by strongly imbricate and spiral foliation of short ascending branches, caudate branch tips bearing smooth, filamentous gemmae in leaf axils, carinate leaves with distinct spine(s) at the base of the dorsal side of the keel, thin-walled, quadrate alar cells arranged in a scalariform manner, narrowly cylindrical capsules with perforate exostome teeth and reduced endostome, and non-collenchymatous exothecial cells. The multistratose alar cells, smooth filamentous gemmae, and non-collenchymatous exothecial cells, as well as phylogenetic inferences based on rbcL sequences, suggest that the species belongs in the genus Clastobryopsis (Pylaisiadelphaceae).

Indopottia irieandoana sp. nov. (Pottiaceae) from Doi Inthanon, northern Thailand

Abstract Indopottia irieandoana H.Akiyama sp. nov. is reported on the basis of two specimens from Doi Inthanon, northern Thailand, as a second species of a genus recently described from southern India. The lack of a peristome precludes unambiguous familial placement based on morphology, but phylogenetic analyses of sequences from a chloroplast and a mitochondrial locus place the new taxon, with strong support, in the Pottiaceae. Its epiphytic habitat on a tree trunk and a branch near the crown of an emergent tree and a strangler tree, respectively, along with morphological features such as spatulate leaves, a single dorsal stereid in the leaf costa, smooth laminal cells, long rostrate opercula, and eperistomate, upright capsules point to a close relationship to Indopottia zanderi. The morphology is compared with that of related genera. A key to similar genera and species is provided.

The New Genus Benitotania (Daltoniaceae, Bryopsida) from Mt. Kinabalu

Abstract A new monospecific genus, Benitotania, is described on the basis of B. elimbata H. Akiyama, T. Yamag, & M. Suleiman, collected in the northern part of Sabah, Malaysia. It appears to be most closely related to Adelothecium and Bryobrothera.

The Climacium-type branch development in the Bartramiaceae and its taxonomic significance

The Bartramiaceae have been defined by unique apple-shaped capsules. The present study deals with the branch development pattern of each genus of the Bartramiaceae to know if the family is consistent in the feature. As a result, it appears that all genera have the Climacium-type of branch development except Catoscopium. This supports the opinion that the Bartramiaceae are monophyletic and also suggests a remote relationship of Catoscopium to the other members of the Bartramiaceae. The Bartramiaceae have long been classified under the suborder Bryineae. The other families of the Bryineae, however, show Bryum-type branch development except the Tim- miaceae. This might indicate that both the Bartramiaceae and Timmiaceae should be separated from the Bryineae.

Morphology and Anatomy of the Peristome in Dozya japonica (Musci) and Reconsideration of Its Affinity to the Leucodontaceae

The most distinctive features in sporophytes ofDozya japonica Lac. are the very thick capsule walls and the presence of parastomes around exostomes. Though D. japonica has no endostomes and its taxonomical position is not clear, the presence of parastomes and the very thick capsule walls indicate no close relationship to Leucodon. Dozya japonica should be included in a new subfamily, Dozyoideae (Leucodontaceae). The genus Dozya is monotypic, with a single species D. japonica Lac. Although the species is widely distributed in East Asia (Japan, Korea, China), it has been collected rarely, except in Japan. The species is restricted to the temperate zone, and is usually found growing on tree trunks and rocks in Japan. The systematic position of Dozya japonica has been considered obscure (Noguchi 1947), and basic

Phylogenetic study of the genus Aptychella (Pylaisiadelphaceae, Musci)

Abstract The relationships among species of the epiphytic genus Aptychella (Broth.) Herzog and Clastobryopsis M. Fleisch. were inferred based on phylogenetic analyses of plastid (rpl16, rps4, and trnL-F) and mitochondrial (nad5) gene sequences as well as morphological features. One of the Asian members, Aptychella robusta (M. Fleisch.) M. Fleisch., exhibits much genetic variation and differentiation among populations, which may reflect the presence of cryptic species. The distinctiveness of Asian members characterized by unicostate leaves, A. brevinervis (M. Fleisch.) M. Fleisch., A. pseudobrevinervis H. Akiyama and A. oblongifolia H. Akiyama, are confirmed. On the contrary, Aptychella proligera, the sole member of the genus known from South and Central America, is shown to have a close relationship to the A. brevinervis complex and preserves low genetic variation among samples collected from geographically remote localities. This lack of variation might be caused by a recent immigration from Asian populations. The Asiatic genus Clastobryopsis is treated as a synonym of Aptychella based on the present phylogenetic analyses. One new combination is proposed—Aptychella imbricata (H. Akiyama et al.) H. Akiyama comb. nov.

Yakushimabryum longissimum (Pylaisiadelphaceae) gen. & sp. nov., from the Yakushima Island, Japan

Abstract Yakushimabryum longissimum gen. & sp. nov. (Pylaisiadelphaceae, Musci), is described from Yakushima Island, Japan, on the basis of morphology and rbcL data. It is characterized by more or less glossy, yellowish green living plants, naked dormant branch buds, absence of costa in the leaves, smooth and linear leaf lamina cells, alar groups composed of a number of quadrate cells arranged in a scalariform manner, ascending branches that are often elongate into slender tips, and filamentous gemmae consisting of smooth, long-rectangular cells. Molecular phylogenetic analyses suggest an affinity neither to Clastobryopsis nor Gammiella, both of which grow in similar habitats and share alar morphology and shape of propagules, but to Isocladiella and Isopterygium, both of which have filamentous pseudoparaphyllia and different types of propagules. A key to the Japanese genera of Pylaisiadelphaceae is provided.