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A Taxonomic Revision of the F Amil Y Ditrichaceae (Musci) of Japan, Korea

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A Taxonomic Revision of the F Amil Y Ditrichaceae (Musci) of Japan, Korea Joum. Hallori Bot. Lab. No. 68: 317- 366 (June 1990) A TAXONOMIC REVISION OF THE F AMILY DITRICHACEAE (MUSCI) OF JAPAN, KOREA AND TAIWAN! 2 2 TOHRU MATSm AND ZENNOSKE IWATSUKI ,3 ABSTRACT. After a taxonomic revision of the family Ditrichaceae of Japan, Korea and Taiwan, there remain 8 genera and 20 species, including one new species (Ditrichum sekil). Cladistic relationships among the genera of the Ditrichaceae were explored. Some taxa were reduced to the synonymy of the species indicated in brackets: Ditrichum subtortile Card. [= D. heteromallum (Hedw.) Britt.], D. divaricatum Mitt. var. exaltatum Card. [= D. divaricalUm Mitt.], and Pleuridium colei (Robins.) Deguchi et Matsui [=Pseudephemerwn nitidum (Hedw.) Hag.]. Detailed descriptions, taxonomic notes, illustrations and distribution maps of each species as well as keys to the genera and species are provided. INTRODUCTION According to Index muscorum japonicorum, Iwatsuki and Noguchi (1973) listed 7 genera and 18 species of Ditrichaceae from Japan. Recently the following two genera were newly found in Japan: Eccremidium Wils. (lwatsuki & Takaki 1979), Pseudephemerum (Lindb.) Hag. (Iwatsuki 1980), and the following three species were added to the moss flo ra of Japan: Ditrichum lineare (Sw.) Lindb. (Matsui et at. 1985), D. rhynchostegium (Matsui & Deguchi 1987) and Pleuridium colei (Robins.) Deguchi et Matsui (Deguchi & Matsui 1985). Matsui and Deguchi (1987) provided a preliminary key to the Japanese species of Ditrichum. In Korea, Choe (1980) listed 4 genera, 6 species and I variety of Ditrichaceae. In Taiwan, Kuo and Chiang (1987) listed 5 genera, 10 species and I variety of the family. Some Asian species of the Ditrichaceae were studied by Seppelt (l982a), Matsui and Deguchi (1987) and Seppelt (1987), and some species were reduced to the synonymy of the species indicated in brackets: Ditrichum dicranelloides Sak. [= D. macrorhynchum Broth. ex Card.], D. kiusiuense Sak. [= D. macrorhynchum Broth. ex Card.] and D. longipes Sak. [= D. pa/lidum (Hedw.) Hampe]. In this revision, we treat all genera and species reported from Japan, Korea and Taiwan. Cladistic analysis of genera of the Ditrichaceae, detailed description and figures will be given. 1 Contribution from the Phytotaxonomical and Geobotanical Laboratory, Hiroshima University, N. Ser. No. 389. 2 Botanical Institute, Faculty of Science, Hiroshima University, Higashi-senda-machi, Hiroshima 730, Japan. 3 Also the Hattori Botanical Laboratory, Obi, Nichinan-shi, Miyazaki-ken 889-25, Japan. 318 10urn. Hattori Bot. Lab. No. 68 1 990 ACKNOWLEDGEMENTS. We wish to express our deep gratitude to Dr. H. Deguchi of Kochi University, Dr. R. D. Seppelt of Antarctic Division who generously gave us invaluable advice. We are also thankful to Dr. H. Robinson of Smith soni an Institute for information on Ditrichum rhynchostegium and to our colleagues of Hiroshima University who kindly placed their specimens at our disposal for the present study. We thank the directors and curators of the following herbaria from which materials including type specimens were supplied on loan: British Museum, London (BM); Duke University, Durham (DUKE); Hattori Botanical Laboratory, Nichinan (NICH); Kyoto University, Kyoto (KYO); Makino Herbarium, Tokyo Metropolitan University, Tokyo (MAK); Museo Botanico, Firenze (FI); National Science Museum, Tokyo (TNS); New York Botanical Garden, New York (NY); Tunghai University, Taichung (TUNG); University of British Columbia, Vancouver (UBC); and Smithsonian Institution, Washington, D. C. (us). Lastly but not least, we heartily thank Dr. G. M. Glime of Michigan Technological University for reading the manuscript. CLADISTIC ANALYSIS OF THE ASIAN DITRICHACEAE The c1adistic relationships of the Asian Ditrichaceae were analyzed at the generic level since homoplasy within a large genus, especially the genus Ditrichum, would not confound the analysis. In this study, character compatibility analysis has been employed. Character compatibility analysis reveals the patterns of agreement and disagreement among cladistic characters and chooses the best characters that reflect evolutionary relationships. Two characters that correctly reflect evolutionary relationships are always compatible, whereas at least one member of an incompatible pair of characters includes parallelisms, reversals or an incorrectly estimated primitive state. Character compatibility analysis finds a mutual compatible character set(s), which is called a clique(s). Meacham (1980, 1981), Estabrook et al. (1977) and Meacham and Estabrook (1985) have written outlines of character compatibility analysis. These analyses were performed by CLIQUE Ver. 2.9 in the program package PHYLIP Ver. 3.0 developed by Dr. J. Felsenstein of Washington University. C1adistic characters To determine the directions of character state trees (CSTs), the Dicranaceae is used as the comparative out-group of the Ditrichaceae. Classification of character states and directions in the Ditrichaceae is discussed below. I. Color of plants: A. Glaucous. B. Not glaucous. CST: B- > A. Only Saelania has glaucous plant color. This color is due to a chemical substance, ( - )-16cx-hydroxykaurane, synthesized by Saelania. In the out-group, the Dicranaceae, no genera have graucous plant color. 2. Leaf arrangement: A. Distichous. B. More than three rows. CST: B-> A. Most genera of the Ditrichaceae and Dicrnaceae have more than three rows. Only Distichium has a clearly distichous leaf arrangement. This character distinguishes Distichium from all other genera in the Ditrichaceae. In the Dicranaceae, no genera have distichous leaf arrangements. 3. Leaf: A. Erect. B. Squarrose. CST: A- > B. T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 319 The leaves of most genera are erect or nearly so. Only Trichodon has squarrose leaves. In the Dicranaceae, some species of Dicranella (e.g. D. palustris), Dichodontium, Oncophorus and others have squarrose leaves, but many genera have erect to erect-spreading leaves. 4. Lamina of leaves: A. Differentiated up to near apex. B. Differentiated only below middle part of the leaf. C. Differentiated only at basal part of the leaf. CST: A- > B-> C. Most genera of the Dicranaceae have a well differentiated lamina that reaches near the apex. But many genera, except Ceratodon and Saelania of the Ditrichaceae, have a poorly differentiated lamina. 5. Cells of sheathing base: A Rectangular to long rectangular. B. Quadrate. C. Linear. CST: C < - A- > B. Ceratodon and Saelania have quadrate cells of sheathing base. Brotherus (1924) recognized this as one of the distinguishing characters of the Ceratodontoideae from other subfamilies of the Ditrichaceae. Distichium has linear cells of sheathing base, which are rare in the Ditrichaceae. Some genera (e.g. Rhabdoweisia) of the Dicranaceae have quadrate cells of sheathing base, but many genera have rectangular to long rectangular cells of sheathing base. 6. Size of cells of sheathing base: A. Normal. B. Large. CST: A- > B. Eccremidium and Pseudephemerum have large cells of sheathing base with thin walls. This character is rare in the Ditrichaceae and the Dicranaceae. 7. Subula of leaves: A. Smooth. B. Mamillose. CST: A- > B. Only Distichium has a mamillose subula in the Ditrichaceae. A few species of Ditrichum (e.g. D. divaricatum) have a mamillose subula, but this character is rare in Ditrichum. In the Dicranaceae, some genera have mamillose or papillose cells of sheathing base, but the numbers are few. 8. Stereid bands. A. Stereid bands differentiated on both abaxial and adaxial sides of guide cells. B. Stereid bands only Differentiated on adaxial size of guide cells. C. Stereid bands not differentiated on either side of guide cells. CST: A- > B- > C. In the Ditrichaceae and the Dicranaceae, many genera have stereid bands on both adaxial and abaxial sides of guide cells. In Pseudephemerum and Trichodon, stereid bands differentiate only on the abaxial side. Eccremidium has no stereid bands. 9. Leaf margin: A. Plane. B. Recurved. CST: A-> B. In the Ditrichaceae and Dicranaceae, most genera have plane leaf margins. Ceratodon, Pseudephemerum and Saelania have recurved margins. In Ditrichum, D. lineare, D. macrorhynchum and some Asian species have recurved leaf margins, but this character is rare in Ditrichum. Ditrichum was treated as having plane margins. 10. Epidermal layer of stem: A. differentiated. B. Not differentiated. CST: A- > B. Epidermal layers of stems are not differentiated in Eccremidium and Pseu­ dephemerum. Lack of this epidermal layer is very rare in the Ditrichaceae and the Dicranaceae. 11. Perichaetialleaf: A. Differentiated. B. Differentiated but very short. C. Not or scarcely differentiated. CST: B< - A- >C. 320 Journ. Hattori Bot. Lab. No. 68 I 990 Most genera in the Ditrichaceae and the Dicranaceae have well differentiated perichaetial leaves. In Ceratodon, perichaetial leaves are well differentiated and very short. This character is very rare in both families. 12. Capsule shape: A. Oblong to cylindric. B. Globose without dehiscent lines. C. Globose with dehiscent lines. CST: A- > B- > C. Globose capsules are found in cleistocarpous genera, such as Eccremidium, Pleuridium and Pseudephemerum. Among these genera, Eccremidium has globose capsules with dehiscent lines. This character distinguishes Eccremidium from all other members of the Ditrichaceae that have globose capsules. 13. Seta length: A. Long. B. Very short. CST: A- >B. Eccremidium, Pleuridium and Pseudephemerum have short setae in the
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