Joum. Hallori Bot. Lab. No. 68: 317- 366 (June 1990)

A TAXONOMIC REVISION OF THE F AMILY (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 ( 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 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 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 : A. Glaucous. B. Not glaucous. CST: B- > A. Only has glaucous 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 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 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 Di­ trichaceae. Other genera in the Ditrichaceae and the Dicranaceae have long setae. 14. Peristome teeth: A. Present. B. Present (outer plate narrower than inner plate). C. Present (peristome teeth differentiated on the special basal tissues. D. Absent. CST: D<- A- >B, A- >C. Most genera in the Ditrichaceae and Dicranaceae have well differentiated

FIG. I . Peristome teeth of various genera of the Ditrichaceae. a. (Deguchi 1307, KOCH). b, (Deguchi 26887, KOCH). c, Ditrichum rhynchrostegium (Deguchi 6807, KOCH). d, Trichodon cylindricus (Roivainen s. n., HIRO). e, Saelania glaucescens (Deguchi 21004, KOCH). Scale bar: 10 jlm. T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 32 1

FIG. 2. Longitudinal section of peristome teeth of various genera of the Ditrichaceae. a, Ceratodon purpureus (Deguchi 13097, KOCH). b, Saelania glaucescens (Iwatsuki 1412, NICH) . c, Distichium capi//aceum (Iwatsuki 1014, NICH). d, Ditrichum rhynchostegium (Deguchi 16630, KOCH) . e, Trichodon cy/indricus (Noguchi 35284, NICH). Scale bars in jlm.

TABLE 1. Cladistic character data. Acronyms correspond to those used in the Fig. 3.

Character States Genus Acronym o 0 0 0 0 000 0 I I I 2 3 45 6 7 8 9 0 2 3 4 5 6

Ceratodon Cer BBAAB AAABA BAABA A Distichium Dis BAABC ABAAA CAAAA A Ditrichum Dit BBACA AAA AA AAAAA A Eccremidium Eec BBACA BACAB CCBDB B Pleuridium Pie BBACA AAAAA CBBDB A Pseudephemerum Pse BBAAA BABBB CBBDB A Saelania Sae ABAAB AAABA CAACA A Trichodon Tri BBBCA AABAA AAAAA A peristome teeth. These are usually papillose throughout (Fig. I). Ceratodon has very unique peristome teeth that are composed of narrow outer plates and wide inner plates (Fig. la). Peristome teeth of Saelania are differentiated on special basal tissue (Fig. 2b). Occurrenses of these characters are very few in the Ditrichaceae and the Dicranaceae. 15. Operculum: A. Present. B. Absent. eST: A-> B. Opercula are present in most genera in the Ditrichaceae and Dicranaceae except Eccremidium, Pleuridium, Pseudephemerum and other cleistocapous genera. 322 Journ. Hattori Bot. Lab. No. 68 I 990

16. Calyptra: A. Long cucullate with smooth surface. B. Mitrate with rough surface. CST: A- > B. Most genera in the Ditrichaceae and the Dicranaceae have cucullate calyptra. Only one genus, Eccremidium, has a mitrate calyptra with rough surface. The data matrix is shown in Table 1.

Results and Discussion Primary analysis using the computer program CLIQUE yielded two largest cliques of 12 mutually compatible characters: Clique I: 1,2,3,5,6,7, 10, 12, 13, 14, 15, 16. Clique II: I, 2, 3,6, 7, 10, 11, 12, 13, 14, 15, 16. These cliques show a branch terminated by Eccremidium and Pseudephemerum and subtended by Pleuridium. To resolve further, a secondary analysis was performed with Eccremidium and Pseudephemerum removed. A single largest clique of 15 characters was found: Clique Ill: 1,2,3,4,5,6, 7, 8, 9, 10, 12, 13, 14, 15, 16. The cladogram defined by the primary and secondary analyses is shown in Fig. 3. In Ditrichaceae, Brotherus (1924) recognized the following three subfamilies: Ditrichoideae, Ceratodontoideae and Distichioideae. Character compatibility analysis suggested that two subgroups can be recognized in the Ditrichaceae of Asia. (I) Ceratodon and Saelania. This group corresponds to subfamily Ceratodontoideae sense Brotherus (1924). (2) Distichium, Ditrichum, Trichodon, Pleuridium, Pseudephemerum and Eccremidium. This group corresponds to subfamilies Ditrichoideae and Distichioideae sensu Brotherus (1924). Cleistocarpous genera (Pleuridium, Pseudephe-

FIG. 3. The c1adogram by the character compatibility analysis. T. MATSul & Z. !WATSUKI: A taxonomic revision of Ditrichaceae of Japan 323 merum and Eccremidium) are separated from other genera in the second group. These cleistocarpous genera plus Ditrichum and Trichodon are a monophyletic group. In this analysis, Brotherus' subfamilies Distichioideae and Ditrichoideae are fused. Based only on Asian genera of the Ditrichaceae, it is difficult to determine whether Distichoideae should be treated as a separate subfamily of the Ditrichaceae.

TAXONOMIC TREATMENT Explanatory notes 1. In the following text, abbreviations of nomenclatural terms follow mainly Wijk et al. (1959); abbreviations of herbarium names follow Holmgren and Keuken (1974). 2. The sizes of cells are given in terms of the size of the lumen in a thoroughly wet condition. We used 90% ethyl alcohol and 2% KOH solution to wet the cells before measurements. 3. Only important illustrations published for Asiatic specimens are listed under "Illustrations" .

Ditrichaceae Limpr., nom. Jam. cons. Laubm. Deutchl. I: 482 (1887). Type genus: Ditrichum Hampe. Distichiaceae Schimp., Syn. Muse. Eur.: 135 (1860), nom. Jam. rejic. Ceratodontaceae Schimp., Syn. Muse. Eur.: 138 (1860), nom.fam. rejic. Plants small, gregarious or loosely tufted, occasionally densely caespistose. Stems erect, sometimes branched, central strands present. Leaves mostly lanceolate or tapering to the apex from oblong base; costa single, usually well developed, occupying most part of subula; cells smooth or rarely roughened, subquadrate to linear, not differentiated at the basal angles. Autoicous or dioicous. Capsules usually oblong to cylindric, erect to inclined, symmetric or asymmetric, sometimes globose; seta usually long, rarely short, straight or arcuate, reddish-brown to bright yellow; exothecial cells mostly rectagular to long hexagonal or quadrate; operculum and annulus usually differentiated, sometimes absent; peristome teeth usually present, 16, split nearly to the base, papillose throughout. Calyptra usually long cucullate, rarely mitrate. Spores usually papillose.

KEY TO THE GENERA 1. Capsules globose; cleistocarpous, operculum not differentiated ...... 2 1. Capsules ovoid-cylindric to cylindric; operculum differentiated ...... 4 2. Capsules dehiscent at middle; stereid bands of leaf costa not differentiated ...... 8. Eccremidium (p. 360) 2. Capsules dehiscent irregularly; stereid bands weakly or clearly differentiated ...... 3 3. Epidermal cells of stems not differentiated; stereid bands of leaf costa weakly differentiated ...... 7. Pseudephemerum (p. 358) 3. Epidermal cells of stems differentiated; stereid bands clearly differentiated ...... 6. Pleuridium (p. 355) 4. Leaves clearly distichous ...... 3. Distichium (p. 330) 4. Leaves in more than three rows ...... 5 5. Plants bluish glaucous...... 2. Saelania (p. 328) 5. Plants not bluish-glaucous ...... 6 324 Journ. Hattori Bot. Lab. No. 68 I 990

6. Leaves contorted when dry; operculum short conic; capsules inclined, strongly furrowed when dry...... 1. Ceratodon (p. 324) 6. Leaves not contorted when dry; operculum high conic; capsules not inclined, smooth when dry (except Ditrichum pallidum) ...... 7 7. Leaves squarrose when dry; peristome teeth incurved when dry .. .. 5. Trichodon (p. 354) 7. Leaves not squarrose when dry; peristome teeth straight or twisted when dry ...... 4. Ditrichum (p. 332)

1. Ceratodon Brid. Bryo!. Univ. I: 480 (l826). Type species: Dicranum purpureum Hedw. [= CeralOdon purpureus (Hedw.) Brid.]. Ceratodon is characterized by the peristome teeth which are composed of a wide inner plate and narrow outer plate, strongly furrowed asymmetric capsules, lanceolate leaves and quadrate cells of sheathing base. Ceratodon has gametophytes similar to Sae/ania, which is a member of Ceratodontoideae sensu Brotherus (1924), but Ceratodon is distinguished from Sae/ania by non-glaucous habit, contorted leaves when dry and small perichaetialleaves.

I. Ceratodon purpureus (Hedw.) Brid. (Fig. 4, Map la) Bryo!. Univ. I: 480 (1826). Dicranumpurpurem Hedw., Spec. Musc.: 136 (l801) ~ Didymodon purpureus (Hedw.) Hook. et Tay!., Musc. Brit.: 65 (1818) ~ Trichostomum purpureum (Hedw.) De Not., Syl!.: 189 (1838). Dicranum purpurascens Hedw., Spec. Musc.: 137 (l801) ~ Ceratodon purpureus (Hedw.) Brid. var. purpurascens (Hedw.) Brid., Bryo!. Univ. I: 483 (1826) ~ Ceratodon purpurascens (Hedw.) Jenn, Man. Moss. W. Pennsylv.: 57 (1913). Dicranum celsii Hedw., Spec. Musc.: 149 (1801 ) ~ Ceratodon purpureus (Hedw.) Brid. var. celsii (Hedw.) Brid., Bryol. Univ. I: 483 (I 826}. Dicranum intermedium Hedw., Spec. Musc.: 138 (1801) ~ Ceratodon purpureus (Hedw.) Brid. var. intermedius (Hedw.) Brid ., Bryo!. Univ. 1: 484 (1826). Bry um biparlitus Dicks. ex With., Syst. Arr. Brit. PI. ed. 4, 3: 818 (1081) ~ Ceratodon purpureus (Hedw.) Brid. var. biparlilus (With.) Brid., Bryol. Univ. I: 484 (1826). Bryum papil/osum Dicks., PI . Crypt. Brit. fasc. 4: 12 (1801) ~ Trichostomum papil/osum (Dicks.) Srn., FI. Brit. 3: 1238 (1804) ~ Didymodon papil/osus (Dicks.) Brid., Spec. Musc. I: 150 (l806) ­ Ceratodon purpureus (Hedw.) Brid . var. papil/osus (Dicks.) Bird., Bryol. Univ. 1: 485 (1826). Bryum slriclum Dicks., PI. Crypt. Brit. fasc. 4: 13 (1801) - Dicranum strictum (Dicks.) Srn., FI. Brit. 3: 1218 (1 804). Dicranum paluslre Brid. ex Schum., Enum. PI . Sael!. 2: 59 (1803) ~ Ceralodon purpureus (Hedw.) Brid. var. paluslris (Schum.) Brid., Bryol. Univ. 1: 485 (1826). Tortula saussuriana P. Beauv ., Prodr.: 93 (l805) ~ Barbula saussuriana (P. Beauv.) Brid., Mant. Musc.: 94 (1819) ~ CeralOdon purpureus (Hedw.) Brid. var. saussuriana (P. Beauv.) Brid., Bryol. Univ. l : 486 (1826). Dicranum erythropum Lam. et Cand., FI. Franc. ed. 2, 2: 477 (l805) ~ Ceralodon purpureus (Hedw.) Brid. var. erythropus (Lam. et Cand.) Lewq., Mem. Soc. Sci. Neuchatel3: 17 (1846). Dicranum longisetum Brid., Mant. Musc.: 66 (1819), horn . illeg. non. D. longisetum Hook., PI. Crypt. Plag. Orb. Nov. Aequin.: 3a (1816). Ceratodonjavanicus Dozy et Molk., PI. Jungh. 3: 337 (1854). Ceralodon purpureus (Hedw.) Brid. var. arislatus Aust., Musci Appal.: 22 (1870). Ceralodon brasiliensis Hampe, Yid. Medd. Nat. For. Kjoebenh. ser. 3, 4: 39 (1872). Ceratodoll capensis Schimp. in Jaeg., Ber. S. Gall. Naturw. Ges. 1870-71 : 460 (1872), nom. inval. in syn. T. MATSUJ & Z. IWATSUKJ: A taxonomic revision of Ditrichaceae of Japan 325

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n 1-20-i iffWhWg FIG. 4. Ceralodon purpureus (Hedw.) Brid. a, plant. b-f, leaves. g, cells at leaf base. h, cells at middle part of sheathing base. i, cells at upper part of sheathing base. j- m, cross-sections of leaf. n, cross-section of stem. 0, perichaetial leaf. p-q, capsules. r, operculum. s, peristome tooth. t, annulus. u, exothecial cells. v, stoma. w, spores. x. calyptra. Drawn from Deguchi 26887 (K OCH). Scale bars in /lm.

Ceratodon condensatus Schimp. in Jang., Ber. S. Gall. Naturw. Ges. 1870-71 : 462 (1872), nom. nudo Ceratodon kerguelensis C. MueJ\., Forshcungesr. Cagelle Bot. 4: 3 (1889), nom. nudo CeralOdon heterophyllus Kindb., Ottawa Net. 5: 179 (1892). Ceratodon microcarpus C. Muel!., Flora 82: 449 (1896). 326 Journ. Hattori Bot. Lab. No. 68 I 9 90

Ceratodon columbiae Kindb., Rev. Bryo!. 23: 20 (1896). Ceratodon sinesis C. Muel!., Nuov. Giorn. Bot. Ita!. n. ser. 3: 104 (1896). CeralOdon conicus C. Muell., Hedwigia 38: 98 (1899), homo iIIeg. non. C. conicus (Hampe) Lindb., Musci Scand .: 27 (1879). Ceratodon brevifolius Mild. ex Par., Ind. Bryo!. Suppl.: lOO (1900), nom. inval. Ceratodon vialis Stir!., Ann. Scott. Nat. Hist. 16: 105 (1905). Ceratodon crassinervis Amann., FI. Mouss. Suisse 2: 379 (1913), homo iIIeg. nOli. C. crassinervis Lor., Bo!. Zeit. 24: 187 (1866). Ceratodon mol/is Amann., Bull. Murithienne 40: 44 (1919). Ceratodon stenocarpous auct. non. 8. S. G.: Noguchi, J. Hattori Bot. Lab. 5: 40 (1951); Chuang, J. Hattori Bot. Lab. 37: 446 (1973). Plants small to medium size, yellow-green to green, often in dense tufts. Stems 4-30 mm high, yellow to reddish brown, sometimes branched, frequently tomentose below, central strands present. Leaves erect, contorted when dry, spreading when wet, ovate to lanceolate, 1.2- 2.5 mm long, apex acute to short-acuminate, margins narrowly revolute, unistratose, entire throughout; costa percurrent to short-excurrent, sometimes ceasing just below the apex, in cross-section stereid bands differentiated on adaxial and abaxial sides of guide cells; cells of sheathing base quadrate to short-rectangular, incrassate, (3- )6-9 J.lm, basal cells generally rectangular, incrassate to thin-walled, l 4-36J.lm. Dioicous. Perichaetialleaves short acuminate from oblong sheathing base. Capsules inclined, sometimes nearly erect, oblong-cylindric, curved, I .5- 2.5 mm long, strongly furrowed when dry, strumose at apophysis; seta 1.5- 2.5 cm long, reddish to dark purple; peristome teeth 16, lanceolate, divided to near the base, outer plate narrow, inner plate wide, each division with filiform segments with weak nodes above, slightly papillose, hyaline above; operculum conic, ca. 0.5 mm long; annulus 2- 3 rows. Calyptra cucullate, ca. 3 mm long. Spores smooth to nearly so, 12- 16J.lm in diam. Chromosome number: n = 13 (Shimotomai & Kimura 1937). Illustration: Noguchi 1987: 121 , Fig. 468. Representative specimens examined: JApAN. HOKKAlOO. Is!. Rishiri, Harada 5342 (HlRO); HONSHU. Aomori-ken, Mt. Osore, Nakajima 21220 (NICH); Akita-ken, Takanosu, Misawa s.n. (NICH); Kanagawa-ken, Ofuna , Hisauchi s.n. (NICH); Tokyo-to, Tokyo Metropolitan University, Inoue s.n. (NICH); Saitama-ken, Mt. Mae-shiro-iwa, Noguchi 35305 (NICH); Niigata-ken, Isl. Sado, Isaka 177 (NICH); Nagano-ken, Mt. Ontake, Deguchi 13097 (KOCH); Yamagata-ken, Mt. Katta, Sato 24 (NICH); Yamanashi-ken, Mt. Fuji, Togashi s.n. (NICH); Fukushima-ken, Hinoemata-mura, Baba 4 (NICH); Gunma-ken, Mt. Haruna, Ohba Sag-203 (NICH); Gifu-ken, Mt. Norikura, Hattori 21405 (NICH); Fukui-ken, Ohno-shi, Sakai 1271 (NICH); Mie-ken, Ohsugi-dani, Magohuku 44 (NICH); Nara-ken, Mt. Ohdaigahara, Mizutani 1629 (NICH); Osaka-fu, Sumiyoshi, Sonobe 9 (NICH); Hyogo-ken, Himeji-shi, Nakajima 9331 (NICH); Tottori-ken, Mt. Daisen, Noguchi 3115 (NICH); Okayama-ken, Mt. Takatsuma, Kiguchi 1824 (NICH); Hiroshima-ken, Hiroshima-shi, Matsui 2111 (HIRO). SHIKOKU. Tokushima-ken, Mt. Tsurugi, Deguchi 21201 (KOCH); Kochi-ken, Mt. Kuishi, Kishi 1277 (KOCH). KyuSHU . Ohita-ken, Aono-dani, Amano 8668 (NICH); Kumamoto-ken, Akamizu, Imae S.n. (NICH) . Fertility: Among 299 specimens examined, 245 are fertile. Ecological notes: Among 165 specimens with ecological data, 77 were collected on soil, 46 on rocks (inc!. 8 on stone walls, two on serpentinite, one on limestome), 19 on sandy soil, 10 on straw roofs, 7 on rotten logs, 5 on concrete, and one on roofing tiles. Distribution: Japan (Hokkaido, Honshu, Shikoku, Kyushu, and Ryukyu), Taiwan; cosmopolitan. Ceratodon purpureus is a very common species that is often troublesome to identify, since this polymorphic species has many described form s and varieties. In the Japanese material, a few specimens have the distinctive short, ovate to oblong-lanceolate, concave T. MATSUI & Z. IWATSUKI : A taxonomic revision of Ditrichaceae of Japan 327

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2. Saelania Lindb. Utkasat Nat. Grupp. Eur. Bladmoss.: 35 (1878). Type species: Bryum caesium ViiI. ex P. Beauv. [ = Saelania glaucescens (Hedw.) Broth.]. Saelania is characterized by the bluish color of plants, narrowly lanceolate leaves, and subquadrate to short rectangular cells of sheathing base. Selania and Ditrichum have similar peristome teeth, but the peristome teeth of Saelania are composed of spe­ cial basal tissues of capsules.

1. Saelania glaucescens (Hedw.) Broth. in Bomanss et Broth. (Fig. 6, Map 2c) Herb. Mus. Fenn. 2: 53 (1884). Trichostomum glaucescens Hedw., Spec. Muse.: 11 2 (180 1) - Leptotrichum glaucescens (Hedw.) Hampe in Schimp., Syn. Muse. Eur.: 146 (1860) - Dilrichum glaucescens (Hedw.) Hampe, Flora 50: 182 (1867). Didymodon aeruginosus Hook. in Brid., Bryol. Univ. I: 516 (1826). Bryum caecium Vill. ex P. Beauv., Prodr.: 45 (1805) - Saeiania caecia (P. Beauv.) Lindb., Utkast Nat. Grupp. Eur. Bladmoss.: 35 (1878). Leptotrichum pruinosum C. Muell., Nuov. Giorn. Bot. Ital. n. ser. 3: 97 (1896) - Saeiania purinosa (c. Muell.) Broth., Nat. Pfl. I: 300 (1901). Leptotrichum subglaucescens C. Muell., Nuov. Giorn. Bot. Ital. n. ser. 3: 97 (1896) - Sealania subglaucescens (C. Muell.) Broth., Nat. Pfl. I: 300 (1901). Plants small, tufted, usually bluish glaucous. Sterns up to 20 mm high, in cross-section composed of thick walled cortical layer, mid-cell layer and thin walled central strands. Leaves T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 329

d e

b ~ FIG. 6. Sae/ania g/aucescens (Hedw.) Broth. in Bomanss et Broth. a, plant. b--e , leaves. f, cells at leaf base. g, cells at middle part of sheathing base (dorsal veiw). h, cells at middle part of sheathing base (ventral veiw). i, cells at middle part of costta (dorsal veiw). j, cells at leaf apex. k-n, cross-sections ofleaf. 0, cross-section of stem. p, perichaetial leaf. q- r, capsules. s, operculum. t, peristome teeth. u, annulus. v, exothecial cells. w, stoma .. x, spores. y, calyptra. Drawn from Deguchi 16432 (KOCH) . Scale bars in jlm . somewhat contorted when dry, erect or erect-spreading when moist, narrowly lanceolate, gradually acuminate, 1-4 mm long, margins irregularly serrate in the upper half, usually reftexed at least below the middle; costa percurrent or shortly excurrent, prominent and toothed at back; upper sheathing basal cells subquadrate or short-rectangular, 8- 24 J.lm long, unistratose along 330 Journ. Hattori Bot. Lab. No. 68 I 990 the margins and sometimes with occasional cell projection as a tooth, lower cells short-rectangular or oblong, 15- 56/lm long. Autoicous. Perichaetial leaves not well differentiated. Capsules exserted, cylindric, erect and symmetric, 1.5- 2.0 mm long, somewhat plicate when dry and empty; seta 3-10 mm long; peristome teeth 16, inserted near the mouth, 0.3- 1.0mm long, split nearly to the base into 2 filiform segments, densely papillose; operculum rostrate, 0.7-0.8 mm long; annulus of 2- 3 rows. Calyptra long cucullate, ca. 2 mm long. Spores somewhat rough, 14-18 /lm in diam. Chromosome number: n= 12+ m (Kumar & Verma 1976),13 (Steere 1954). Illustration: Noguchi 1987: 121, Fig. 46A. Representative specimens examined: JAPAN. HOKKAlDO. Hidaka-shicho, Mt. Apoi, Shimizu s.n. (NICH). HONSHU. Aomori-ken, Mt. Hakkohda, Deguchi 16432 (KOCH); Nagano-ken, Mt. Yatsugatake, Furuki 3601 (HIRO); Yamanashi-ken, Mt. Fuji, Watanabe 22169 (NICH~; Fukushima-ken, Minami-aizu-gun, Higuchi 10434 (NI CH); Hyogo-ken, Mt. Rokkoh, Matsui2011 (HIRO); Tottori-ken, Mt. Daisen, Koponen 21854 (NICH). Fertility: Among 61 specimens examined, 51 are fertile. Ecological notes: Among 51 specimens with ecological data, 35 were collected on soil, and IS on rocks (inc!. 3 on serpentinite). Distribution: Japan (Hokkaido and Honshu), Asia, Europe, Greenland, North America, South America, southern Africa and New Zealand. Saelania glaucescens can be easily recognized by its bluish color. Such characteristic coloration of plant has been erroneously understood to be mycelia covering the plants. However, it is not mycelia, but a chemical substance, (-)- 16a-hydroxykaurane, produced by Saelania glaucescens (Nilsson & Martensson 1971; Martensson & Nilsson 1974). We confirmed that the mycelia-like substance on leaves disappeared in acetone, and white, acicular crystals were left on a glass slide after evaporation of solvent.

3. Distichium B. S. G., nom. cons. Bryo!. Eur. 2: 153 (1846). Type Species: Cynontodium capillaceum Hedw. [=Distichium capillaceum (Hedw.) B.S.G .]. Cynontodium Hedw., Spec. Musc.: 57 (1801), nom. rejic. Swartzia Brid., J. f. Bot. 1800: 289 (1801), homo illeg., non. S wartzia Schreb., Gen Syst. 2: 518 (1791) (Phan.). Cynodontium Brid., Musc. Recent. Sup!. 1: IS (1806), nom. rejic. Distichium is characterized by the distichous leaf arrangement and mamillose subula.

1. Distichium capillaceum (Hedw.) B. S. G. (Fig. 7, Map 2a) Bryo!. Eur. 2: 153 (1846). Cynontodium capillaceum Hedw., Spec. Musc.: 57 (1801) - Swartzia capillacea (Hedw.) Brid., J. f. Bot. 1800: 289 (1801) - Cynodontium capillaceum (Hedw.) Brid., Musc. Recent. Suppl. I: 158 (1806). Swartzia montana Lindb., Act. Soc. Sci. Fenn. 10: 16 (1871), nom. illeg. incl. spec. prior. Distichium montanum Hag., K. Norsk. Yid . Selsk. Skrift 1910: 61 (1910), nom. iIleg. incl. spec. p;ior. Distichium setifolium C. Mue1!. , Linnaea 43: 396 (1882). Plants slender, forming dense tufts, green above, brown below. Stems 10-40 mm high, erect, densely tomentose below, nearly square in cross-section, central strands present, epidermal layer 2- 3 rows, cells smaller, incrassate; leaves distichous, appressed, 3- 5 mm long, subula mamillose, apices acute, occasionally weakly toothed, base clasping stem, margins weakly convolute below; costa occupying most part of subula, in cross-section stereid bands differentiated on both ventral and dorsal sides of guide cells; upper sheathing basal cells quadrate to rectangular, 3-15/lm T. MATSUI & Z. [WATSUKl: A taxonomic revision of Ditrichaceae of Japan 331 )oUt ~ ) r ) "

l .• ~(r 0 (( 0 I 0 1-20-4 0 0 ( ( I 0 0 -t' ,') 0 (( " Q ( ) 0 l '"(". ./ 0 .0 ~ 1: I ] ·

FIG. 7. Distichium capillaceum (Hedw.) B. S. G. a, plant. b-e, leaves. f- g, cells at leaf base. h, cells at middle part of sheathing base. i, cells at upper part of sheathing base. j, middle part of subula (dorsal view). k, leaf apex. l-q, cross-sections ofleaf. r, cross- section of stem. s, perichaetial leaf. t- u, capsules. v, peristome teeth. w, exothecial cells. x, stoma. y, calyptra. Drawn from Deguchi 26887 (KOCH). Scale bars in jjm.

long, smooth, basal cells long-rectangular to linear, 3~75)lm long. Autoicous. Perichaetial leaves scarcelly differentiated. Capsules erect, symmetric, oblong to cylindric, ca. I mm long, sometimes furrowed; seta erect, ca. 10 mm long; peristome teeth 0.1--0.2 mm long, divided to near the base, lightly papillose reddish yellow; operculum conic, ca. 0.3 mm long; annulus of 2 rows. Calyptra cucullate, I mm long. Spores nearly smooth, 16--22)lm in diam. Chromosome 332 10urn. Hattori Bot. Lab. No. 68 I 990

number: n= 14 (Smith & Newton 1968),28 (Anderson & Crum 1958). lIlustration: Noguchi 1987: 123, Fig. 47. Representative specimens examined: JAPAN. HOKKAlDO. Is!. Ri shiri, Shimizu s. n. (NICH) . HONSHU. Nagano-ken, Mt. Yatsugatake, Mizutani s. n. (NICH); Yamanashi-ken, Mt. Kitadake, Osada 74006 (NICH); Saitama-ken, Jyumonji pass, Deguchi 26886 (KOCH); Hyogo-ken, Mt. Funakoshi, Tatebe 2136 (NICH). SHIKOKU . Tokushima-ken, Mt. Tsurugi, Deguchi 20093 (KOCH). Fertility: Among 52 specimens examined, 29 are fertile. Ecological notes: Among 45 specimens with ecological data, 42 were collected on rocks (inc!. 21 on limestone), and three on soil. Distribution: Japan (Hokkaido, Honshu and Shikoku), northern and central Asia, Europe, North and South Americas, New Guinea, Australia and New Zealand. Distichium capillaceum has a clearly distichous leaf arrangement. The scabrous subula is also a useful character for the recognition of the species, but the gametophyte characters, except the distichous leaf arrangement, are very similar to those of Ditrichum divaricatum. Sainsbury (1955) and Scott and Stone (1976) mentioned that the general appearance and habit were the same as in Ditrichum. In addition, these two genera sometimes share the same habitat.

4. Ditrichum Hampe, nom. cons. Flora 50: 181 (1867). Type species: Trichostomum homomallum Hedw. [= Dilrichum heleromallum (Hedw.) Britt.). Leptotrichum Hampe, Linnaea 20: 74 (1847), horn. illeg., non Leptotrichum Corda, Icon. Fung. 5: 10 (1842) (Fungi). Plants loosely to densely tufted, yellowish green to dark green. Stems 2-70 mm high, simple or branched, with central strands. Leaves erect to erect-patent, lanceolate, ovate-lanceolate or oblong-ovate, with sheathing base, mostly subulate, sometimes secund; costa occupying most part of subula, sheathing basal narrow above and sometimes absent above the shoulders; cells at upper of shating base subquardrate to linear; lower cells oblong to linear; stereid bands, differentiated on both sides of guide cells or only on abaxial side. Autoicous or dioicous. Capsules erect or inclined, cylindric or oblong, often asymmetric, rarely furrowed when dry; setae straight; peristome teeth 16, usually split to the base into two filiform divisions, occasionally imperfectly split, papillose throughout; exothecial cells rectangular; annulus of two to three rows; stomata present at apophysis; operculum high conic. Calyptra cucullate. Spores papillose. Ditrichum is characterized by the filiform and papillose peristome teeth.

KEY TO THE SPECIES I. Subula of leaves mamillose; leaves with elongate sheathing base; cells at upper part of sheathing base irregularly quadrate, incrassate; cells at middle to base of sheathing base linear, with thin wall ...... 11. D. divaricatum (p. 352) I. Subula smooth or nearly so; leaves without elongate sheathing base; cells at upper part of sheathing base quadrate or rectangular; cells at middle to base of sheathing base quadrate or rectangular moderately thickened walls ...... 2 2. Plants large, stems to 7 cm long; basal leaf cells with nodulosely thickened longitudinal walls ...... 6. D. crispatissimum (p. 341) 2. Plants small, stems usually 2 cm long; basal leaf cells with evenly thickened longitudinal walls...... 3 T . MATSUI & Z. (WATSUKI : A taxonomic revision of Ditrichaceae of Japan 333

3. Leaf margins recurved ...... 4 3. Leaf margins plane ...... 5 4. Leaves short, ca. I mm long, appressed when dry ...... 2. D. lineare (p. 336) 4. Leaves long, usually \.5- 2 mm long, erect spreading when dry ...... 3. D. macrorhynchum (p. 337) 5. Leaves triangular; middle part of sheathing base bistratose ...... 6 5. Leaves from oblong-ovate base gradually narrowed into subula; middle part of sheathing base unistratose ...... 7 6. Leaves erect to erect-spreading, \.5- 3.0mm long...... 5. D. brevidens (p. 340) 6. Leaves appressed to the stems, 0.8- 1.6 mm long ...... 4. D. zonatum (p. 339) 7. Cells of sheathing base quadrate; crose-sections of subula biconvex .. 7. D. sekii (p. 344) 7. Cells of sheathing base rectangular (-linear); cross-sections of subula not biconvex. . . . 8 8. Dioicous. Capsules erect, oblong to cylindric, symmetric; seta 10-25 mm long ...... I. D . heteromallum (p. 333) 8. Autoicous. Capsules erect to inclined, cylindric, weakly asymmetric; seta (10-)20-40 mm long ...... 9 9. Perichaetial leaves without sheathing base, 2- 3 mm long; costa occupying most of upper part of sheating base; spores finely papillose, 16-25/lm in diam ... 8. D. pallidum (p. 344) 9. Perichaetial leaves with long sheathing base, 4-5 mm long; costa occupying nearly 1/2 of the upper part of sheathing base; spores usually reticulate, 12- 17/lm in diam ...... 10 10. Spores finely papillose; seta bright yellow ...... 9. D. difficile (p. 346) 10. Spores vermiculately and striately papillose; seta usuall y reddish brown ...... 10. D. rhynchostegium (p. 349)

1. Ditrichum heteromallum (Hedw.) Britt. (Fig. 8, Map 2b) N. Amer. FI. 15: 64 (1913). Weissia heteromalla Hedw., Spec. Muse.: 71 (180l). Didymodon homomallum Hedw., Spec. Muse.: 105 (1801) - Trichostomum homomallum (Hedw.) B.S.G., Bryo!. Eur. 2: 130 (1 843) - Leplotrichum homomallum (Hedw.) Hampe in C. Muel!., Syn. Muse. 1: 453 (1848) - Ditrichum homomallum (Hedw.) Hampe, Flora 50: 182 (1867). Ditrichum subtortile Card., Bull. Herb. Boiss. ser. 2, 7: 716 (1907), syn. novo Plants small, loosely to densely tufted, yellowish green. Stems 3- lOmm high, simple, sometimes branched, with central strands. Leaves \.5- 3.0 mm long, erect to spreading, slightly secund, from ovate to oblong, gradually tapering to long channelled subula; margins plane throughout, unistratose below, bistratose in middle to upper parts; costa broad, occupying most of subula, in cross-section stereid bands distinct on dorsal side, poorly developed on ventral side of median guide cells; upper cells of sheathing base rectangular to linear, juxtacostal cells more elongated, basal cells elongate rectangular to linear. Rhizoidal tubers rarely present. Dioicous. Perichaetial leaves with long sheathing base, 2- 3 mm long. Capsules erect, oblong to cylindric, symmetric, urn 0.4-1.3 mm long, reddish brown, smooth even when dry; setae 10-25 mm long, reddish to purple brown; peristome teeth pale orange, ca. 0.3 mm long, lightly papillose throughout: exothecial cells rectangular, usually 10-12/lm wide; annulus of two to three cell-rows; stomata at apophysis; operculum conic. Calyptra cucullate. Spores lightly papillose, 10-14/lm indiam. Chromosome number: n= 13 (Smith&Newton 1966), l3+m(Kumar& Verma 1979). lliustration: Takaki 1957: 96, Fig. I; Noguchi 1987: 117, Fig. 448. Representative specimens examined: JAPAN. HOKKAIDO. Mt. Soranuma, Notani 36 (NICH). HONSHU. Aomori-ken, Mts. Hakkohda, Deguchi 18635 (KOCH); Nagano-ken, Mt. Ontake, Kuno 24 16 (NICH); 334 Journ. Hattori Bot. Lab. No. 68 1 990

q a sJ V,U (ff).

T o "'J.. T .,.g 1

oT o o... rf{14r 1

1-20-1

c

d T oT g o I 1

FIG . 8. Ditrichum he/eromallum (Hedw.) Brilt. a, plant. b-d, leaves. e, cells at leaf base. f, cells at middle part of sheathing base. g, cells at leaf apex. h- I, cross-sections of leaf. m, cross-section of stem. n, perichaetial leaf. o-p, capsules. q, peristome tooth. r, annulus. s, exothecial cells. t, stoma. u, spores. v, calyptra. w, rhizoidal tuber. a, d , g, i- k, m, n, 0, and q- v drawn from Ma/sui 1118 (KOCH); lH:, e- f, h, I, and p from isotype of D. subtortile (KYO), w from Higuchi 2953 (HIRO) . Scale bars in Ilm.

Gunma-ken, Mt. Shibutsu, Noguchi 33960 (NICH); Yamanashi-ken, Mt. Kitadake, Higuchi 2958 (HIRO); Fukushima-ken, Hinoemata-mura, Higuchi 21031 (NICH). Tochigi-ken, Mt. Nasu, Faurie 119- isotype of D. subtortile (KYO); Ishikawa-ken, Mt. Hakusan, Akiyama 3102 (KYO); Toyama-ken., Mt. Tateyama, Nakajima 1202 (NICH); Nara-ken, Mt. Hidega-take, Deguchi 7846 (KOCH). SHIKOKU. Tokushima-ken, Mt. Tsurugi, Deguchi 18343 (KOCH); Kochi-ken, Mt. Myojin, Deguchi 21481 (KOCH). TAIWAN . M t. Ari, Noguchi s. n. (NICH). T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 335

• Cera todon purpureus • DitrichuII lineare . ~ Ditrichum djffjC ~ _ • .J>1 ' U (f I --- -- cl

• Pleuridium 5ubuJatufIl • Trichodon cyUndricus • PleuridiulJI julaceum

... -:~.~ / :I I I ! 'I I ' I I I I

. ~ old ·~t ~ \ __u i U, is .. . ti

MAP 336 Journ. Hattori Bot. Lab. No. 68 I 990

Fertility: Among 50 specimens examined, 45 are fertile. Ecological notes: Among 32 specimens with ecological data, 26 were collected on soil, and six on rocks. Distribution: Japan (Hokkaido, Honshu and Shikoku), Taiwan, India, Western Himalaya, Europe, and North and South Americas. Ditrichum heteromallum has a general appearance similar to that of D. macrorhynchum, but is easily recognized by the following characters: (I) leaves ovate to oblong, gradually tapering to a long subula, and leaf margins plane and bistratose, (2) elongate rectangular to linear basal cells at sheathing base, and (3) lightly papillose peristome teeth. Cardot (1907) described D. subtortile from Mt. Nasu, central Japan. It is conspecific with D. heteromallum, although the type material of D. subtortile has a shorter seta than the usual from of D. heteromallum. However, even in a single collection, we found that setae are very variable in length. Since we could not find any distinguishing characters between D. subtortile and D. heteromallum, the former should be reduced to synonymy of the latter. The rhizoidal tubers of this species were first discovered by Risse (1985) for European material, and Deguchi and Matsui (1986) found them in Japanese material.

2. Ditrichum lineare (Sw.) Lindb. (Fig. 9, Map Ib) Acta Soc. Sci. Fenn. 10: 108 (1871). Didymodon linearis Sw., Adnot. Bot. : 100 (1829). Trichos/omum vaginans Sull ., Musci Allegh.: 42 (1845) - Lep/olrichum vaginans (Sull. ) Mol., Jahresber. Naturh. Ver. Pasau 10: 109 (1875).

FIG. 9. Ditrichum lineare (Sw.) Lindb. a, plant. b--d, leaves. e, cells at leaf base. f, cells at middle part of sheathing base. g, cells at leaf apex. h- I, cross-sections of leaf. m, cross-section of stem. n, rhizoidal tuber. Drawn from Ma/sui 333 (KOCH). Scale bars in /lm. T. MATSUI & Z. IWATSUKI : A taxonomic revision of Ditrichaceae of Japan 337

LeplOtrichum avimontanum Schimp. in Limpr., Laubm. Deutsch. I : 499 (1887), nom. nudo in synon. Plant small. Stems 2- 7 mm high, nearly triangular in cross-section, central strands differentiated, epidermal cells thick-walled. Leaves 0.5- 1.4 mm long, 0.15--0.35 mm wide, weakly appressed to the stem, imbricate and more or less triseriate, oblong-lanceolate, widest at or little below the middle, gradually narrowed towards the apex, subula short, channelled; leaf margins mostly bistratose, faintly and irregularly recurved above; costa wide, occupying about 1/3 of the width of the leaf base, with a weakly developed stereid band on the abaxial side of the guide cells; cells of sheathing base with smooth, thin to moderately thickened walls, rectangular, 6--8 J1.m wide at the base of sheathing base, becoming slightly shorter and narrower towards the leaf apex. Rhizoidal tubers present. Dioicous. Sporophytes unknown for Japanese plants. Chromosome number: n = 13 + m (Anderson & A I-Aish 1963). Illustration: Matsui et al. 1985: 34, Fig. 1. Representative specimens examined: JAPAN. HONSHU. Chiba-ken, Takahagi, Sasaoka 5362 (TNS); Gifu-ken, Nakatsu-gawa, Hachiya 37 (NICH); Hyogo-ken, Nishinomiya-shi, Matsui 660 (KOCH); Hiroshima-ken, Isl. Miyajima, Matsui 525 (KOCH) . SHIKOKU . Ehime-ken, Mt. Kamega-mori, Marsui 332 (KOCH) . KYUSHU. Kumamoto-ken, Mt. Nekodake, lmae 363 (NICH ). Fertility: All 15 specimens examined are sterile. Ecological notes: Among 13 specimens with ecological data, 12 were collected on soil, and one on sand. Distribution: Japan (Honshu, Shikoku and Kyoshu), Europe and North America. Ditrichum lineare is characterized by the weakly appressed leaves, short subula and recurved leaf margins. Ditrichum lineare is similar to D. pusillum. However, it is distinguished from D. pusi/lum by its rectangular cells of sheathing base.

3. Ditrichum macrorhynchum Broth. ex Card. (Fig. 10, Map 2c) Bull . Soc. Bot. Geneve, ser. 2, 1: 121 (1909). Dit,ichum kiusiuense Sak., Bot. Mag. Tokyo 46: 737 (1932). Dit,icl!um dicranelloides Sak., Bot. Mag. Tokyo 53: 62 (1 939). Plants small, densely tufted, yellowish green to bright green. Stems 7'-20 mm high, simple or sometimes branched, with central strands. Leaves 1- 3 mm long, erect to erect-patent, triangular, gredually tapering to the apex; margins recurved, sometimes unistratose in basal part, bistratose in middle and upper parts; costa short excurrent, in cross-section stereid bands dis­ tinct on dorsal side, poorly developed or rarely absent on the ventral side of guide cells; upper cells of sheathing base rectangular to elongated rectangular, basal cells short rectangular to rec­ tangular. Rhizoidal tubers present. Dioicous. Perichaetial leaves with long sheathing base. Capsules erect, ovate cylindric, symmetric, urn ca. I mm long, reddish brown to dark brown, smooth even when dry; setae 10-20 mm long, reddish to dark brown; peristome teeth orange, divided into two divisions to the base, 0.20--0.35 mm long, densely papillose throughout; exothecial cells rectagular, 14-20 J1.m wide; annulus of three cell rows; stomata at apophysis; operculum conic to elongate conic. Calyptra long cucullate. Spores papillose, 11 - 15 J1.m in diam. Chromosome number unknown. Illustration: Noguchi 1987: 117, Fig. 44C. Representative specimens examined: JAPAN. HONSHU. Miyagi-ken, Sendai-shi, Uyematsu s.n. - paratype of D. macrorhynchum (NICH); Niigata-ken, Kawachi-mura, lkegami 8159 (NICH); Nagano-ken, Mt. Ontake, Noguchi 32118 (NICH); Fukui-ken, Ohno-shi, Sakai 805 (TNS); Wakayama-ken, Honguu-cho, Deguchi 7309 (KOCH); Nara-ken, Higashi-yoshino-mura, Deguchi 8159 (KOCH); Osaka-fu, Nishinose-mura, Nakajima 6988 (NICH); Hyogo-ken, Mt. Myohken, Mizutani 4408 (NICH); Tottori-ken, Mt. Nagi-san, Noguchi 24085 (NICH); Hiroshima-ken, Mt. Jiyama, Malsui 1139 (KOCH). SHIKOKU. Ehime-ken, Mt. Sasaga-mine, Matsui 338 Journ. Hattori Bot. Lab. No. 68 I 990

a t T o 1. f--20-4 k 1

1o o I

b c

To ~ 1

FIG. 10. Dilrichum macrorhynchum Broth. ex Card. a, plant. b-f, leaves. g, cells at leaf base. h, cells at middle part of sheathing base. i, cells at leaf apex. j-n, cross-sections ofleaf. 0 , cross-section of stem. p, rhizoidal tuber. q, perichaetialleaf. r- s, capusules. t, peristome teeth. u, annulus. v, exothecial cells. w, stoma. x, spores. y, calyptra. a, j. 0-<\, and t- x drawn from Deguchi 8159 (KOCH); b-d, g-i, k- n, r- s, and y from paratype of D. macrorhynchum (NICH); e from holotype of D. dicranelloides (MAK); f from holotype of D. kiusiuense (MAK). Scale bars in /lm . T. MATSUI & Z. IWATSUKI : A taxonomic revision of Ditrichaceae of Japan 339

1125 (KOCH); Tokushima-ken, Mt. Tsurugi, Deguchi 19424 (KOCH); Kochi-ken, Mt. Kuishi, Kishi 232 (KOCH). KVUSHU. Fukuoka-ken, Nanbata-mura, Kuwahara 2428 (NICH); Kumamoto-ken, Hitoyoshi, Maebara II- holotype of D. kiusiuense (MAK; isotype NICH); Ohita-ken, Shin-yabakci, Sano 936 (NICH); Miyazaki-ken, Mt. Kuniini-dake, Kuwahara 1435 (NICH); Kagoshima-ken, Shigetomi-mura, Doi s.n. - holotype of D. dicranelloides (MAK ; isotype NICH) . RVUKYU . Is!. Okinawa, Yamashiro 61 (NICH). Fertility: Among 85 specimens examined, 60 are fertile. Ecological notes: Among 55 specimens with ecological data, 42 were collected on soil, 12 on rocks, and one on volcanic ash. Distribution: Japan (Honshu, Shikoku, Kyushu and Ryukyu). Ditrichum macrorhynchum has lanceolate leaves similar to those of D. brevidens, D. linea re and D. zonatum, but it can be easily distinguished from D. lineare and D. zonatum by the longer leaves (1.5- 2.0mm long) that are not appressed to the stem even whey dry. D. macrorhynchum shows great variations in (1) shape of the cells of sheathing base: rectangular to linear, sometimes short rectangular at basal part, (2) cell walls varying from thin to thick, and (3) stereid band developed in various degrees on ventral side. The rhizoidal tubers of this species were discovered by Matsui (1986).

4. Ditrichum zonatum (Brid.) Kindb. (Fig. 11 , Map 1b) Bot. Not. 1883: 146 (1882). Weissia zonata Brid., Bryo!. Univ. 1: 364 (1826) - Ditrichum homomallum (Hedw.) Hampe var. zonatum

b

1-20-1

FIG . 11. Ditrichum zonatum (Brid.) Kindb. a, plant. k , leaves. d, cells at leaf base. e, cells at uppe part of sheathing base. f, cells at leaf apex. g- j, cross-sections of leaf. k, cross-section of stem. Drawn from Diguchi 21046 (KOCH) . Scale bars in }lm. 340 Journ. Hattori Bot. Lab. No. 68 I 990

(Brid.) Lindb., Musci Scand.: 26 (1876) - Ditrichum heteromallum (Hedw.) Britt. var. zona turn (Brid.) Podp., Consp.: 93 (1954). Leptotrichum molendoanurn Lor., Verh. Zool. Bot. Ges. Wien 17: 683 (\867), nom. nudo in synon. LelOtrichum confertum Stirt., Ann. Scott. Nat. Hist. 15: 112 (1906). Plants small, densely tufted, yellowish green to dark green. Stems 10-20mm high, simple or sometimes branched, with central strands. Leaves 0.8- 1.6 mm long, erect, triangular, gradually tapering toward the apex; margins plane throughout, bistratose except basal part; sheathing base unistratose below, bistratose in the upper part; costa short excurrent, broad, in cross-section stereid bands poorly developed on dorsal side and poorly developed or absent on ventral side of guide cells; cells of upper part of sheathing base quadrate to rectagular, basal cells quadrate to short rectangular. Dioicous. Sporophytes unknown. Chromosome number unknown. Specimens examined: JAPAN. HOKKAlDO. Mt. Rishiri, Deguchi 12118 (KOCH); Mt. Yohtei, Deguchi 12647 (KOCH) . HONSHU. Aomori-ken, Mts. Hakkohda, Deguchi 16146 (KOCH); Yamanashi-ken, Mt. Fuji, Deguchi 21046 (KOCH); Nagano-ken, Mt. Tsubakuro, Deguchi 12896 (KOCH); Nara-ken, Mt. Ohdaigahara, Deguchi 7926 (KOCH). Fertility: All 6 specimens examined are sterile. Ecological notes: Five specimens with ecological data were collected on rocks (incl. two on lava). Distribution: Japan (Honshu), Europe, North America. New to Japan. Since sporophyte is unknown, the taxonomic position of Ditrichum zonatum has not been well settled yet. It is often treated as a variety of D. heteromallum. We like to follow Limpricht (1887), Dixon (1924), Lawton (1971), Smith (1978) and others, who treated it as a distinct species. D. zonatum is clearly separated from D. heteromallum by the appressed leaves with bistratose sheathing base, characters not common in the genus Ditrichum.

5. Ditrichum brevidens Nog. (Fig. 12, Map Ib) J. Jap. Bot. 20: 255 (1943). Plants small, loosely to densely tufted, yellowish green. Stems 10-15 mm long, simple, sometimes branched, with central strands. Leaves 1.5- 3.0 mm long, erect to spreading, from oblong to lanceolate, gradully tapering to channelled subula; margins plane throughout; sheathing base unistratose below, bistratose in middle to upper parts; costa broad, occupying most of subula, in cross-section stereid bands distinct on dorsal side, poorly developed on ventral side of guide cells; cells at upper part of sheathing base rectangular to linear, juxtacostal cells more elogated, basal cells elongated rectangular to linear. Dioicous. Perichaetialleaves with long sheathing base, 2- 3 mm long. Capsules erect, oblong to cylindric, symmetric to weakly asymmetric, urn 0.4--1.3 mm long, reddish brown, smooth even when dry; setae 10-25 mm long, reddish to purple brown; peristome teeth pale orange, divided into two divisions to the base, ca. 0.3 mm long, weakly papillose throughout; exothecial cells rectangular, usually 1O- 12jlm wide; annulus of two to three cell-rows; stomata at apophysis; operculum conic. Spores weakly papillose, 10-14 jlm in diam. Chromosome number unknown. Illustration: Noguchi 1943 : 256, Fig. 27 . Representative specimens examined: TAIWAN. Mt. Kodama, Noguchi 6352 - holotype of D. brevidens (NICH); Mt. Ali, Asahina & Ogata s.n., Sasaoka 3886 (TNS) . Fertility: All three specimens examined are fertile. Ecological notes: Ecological data unknown. Distribution: Endemic to Taiwan. T. MATSUl & Z. ]WATSUKI: A taxonomic revision of Ditrichaceae of Japan 341

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0T 0 0 ?~~ru~T I lOO\lD8~[ I \ I~ nn, l8RmRn~

FIG. 12. Ditrichum brevidens Nog. a, plant. H, leaves. f, cells at leaf base. g, cells at middle part of sheathing base. h, cells at leaf apex. i-k, cross-sections of leaf. I, cross-section of stem. m, capsule. n, exothecial cells. 0, stoma. a, c-n, h and m drawn from holotyp of D. brevidens (tns); b, e, f- g, i-I and n-o from Asahina & Ogata S.n. (TNS) . Scale bars in jlm.

Ditrichum brevidens is characterized by leaves from oblong to lanceolate, gradually tapering to channelled subula, bistratose sheathing base and plane leaf margins. Ditrichum brevidens is similar to D. macrorhynchum, but it is easily recognized by the (I) bistratose at moddle to upper parts of sheathing base and (2) plane leaf margins.

6. Ditrichum crispatissimum (c. Muel!.) Par. (Fig. 13, Map lc) Ind. Bryo!. supp!.: 131 (1900). Leptotrichum crispatissimum C. Muel!., Nuov. Giorn. Bot. Ita!. n. ser. J: 97 (1896). Didymodonjiexicaule (Schwaegr.) Roehl var. sterile De Not., Syll .: 198 (1838). Ditrichum giganteum Williams, Bul!. New York Bot. Gard. 2: 113 (1903). Leptotrichum fnfuscatum Stirt., Ann. Scott. Nat. Hist. 46: 112 (1 903). Ditrichumjiexicaule (Schwaegr.) Hampe fo. luxurians Bryhn ex Holz., Bryologist 13: 53 (1910), nom. nudo Leptotrichum jiexicaule (Schwaegr.) Hampe var. longifolium Zett., K. Svensk. Vet. Akad. Hand!. 5: 23 (1865) - Ditrichum jiexicaule (Schwaegr.) Hampe var. longifolium (Zett.) Hag., Tromso Mus. Aarsh. 22: 40 (1899). Ditrichumfragilicuspis Dix. et Sainsb., J. Bot. 71 : 213 (1933). Plants large, dull green to yellowish green above, brown below in fresh material, in soft, 342 Journ. Hattori Bot. Lab. No. 68 I 990

FIG. 13 . Ditrichum crispatissimum (c. MueJ!.) Par. a, plant. b--c, leaves. d, cells at leaf base. e- f, cells at lef margins. g, cells at middle part of sheathing base. h, ceUs at leaf apex. i- m, cross-sections of leaf. n, cross-section of stem. a-m drawn from lectotype of D. crispatissimum (BM); n from Deguchi 17024 (KOCH). Scale bars in /lm. thick tufts. Stems 20-70mm long, simple or rarely branched, with central strands. Leaves faJcate-secund from convolute sheathing base gradually narrowed into long, channelled subula, 6-7 mm long, with entire margins except upper parts where slightly serrulate; costa occupying about 1(3 the width of leaf base, in cross-sections stereid bands poorly developed on both sides of guide cells; cells at upper part of sheathing base rhomboidal to irregular at and near margins and oblong to short vermicular near costa, with evenly thickened walls; basal cells elongate rectangular to vermicular near costa, with evenly thickened walls or thin transverse and thick, nodulose longitudinal walls. Dioicous. Sporotphytes unknown for Japanese plants. Chromosome number unknown. Illustration: Noguchi 1987: 117, Fig. 44A (as D. flexicaule). Representative specimens examined: JAPAN. HOKKAIDO. Shiribeshi-shicho, Mt. Obira, Harada 4699 (HIRO). HONSHU. Nagano-ken, Mt. Ontake, Kuno 2382 (NICH); Saitama-ken, Mt. Bukoh, Noguchi 35138 (NICH) . SHIKOKU. Tokushima-ken, Mt. Tsurugi, Matsui 1103 (KOCH); Kochi-ken, Mt. Ishidata, Deguchi 19294 (KOCH); Ehime-ken, Tengu-kohgen, Kiguchi 6471 (NICH). TAIWAN . Niitaka-shita, Noguchi s.n. (NICH). Fertility: All 44 specimens examined are sterile. Ecological notes: Among 39 specimens with ecological data, 36 were collected on rocks (inc!. 34 on lime tone and one on serpentinite), two on soil, and one on rotten stump. Distribution: Japan (Hokkaido, Honshu and Shikoku), Taiwan, New Guinea, New Zealand, Europe and North and Central Americas. Ditrichum crispatissimum is distinguished from other Ditrichum species by the large plants that are similar to a species of Dicranodontium and the cells at basal part T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 343 of sheathing base with nodulose longitudinal walls. Frisvoll (1985) recognized two different taxa, D. fiexicaule (Schwaegr.) Hampe and D. crispatissimum (c. Muel!.) Par. He distinguished these from each other by 16

T g on 1 v T o o b o... 1 d c a T o To o :;; I 1

FIG. 14. Ditrichum sekii Ando et Deguchi ex Matsui et Iwats. a, plant. b-e, leaves. f, cells at leaf base. g, cells at upper part of sheathing base. h, cells at margins of subula. i, cells at leaf apex. j-l, cross-sections of leaf. m, cross-section of stem. n-o, capsules. p, operculum. q, peristome teeth. r, exothecial cells. s, stoma. t, spores. u, calyptra. v, perichaetialleaf. a-o, q- t and v drawn from holotype of D. sekii (HIRO); p and u from Mayebara s.n. (NICH). Scale bars in /lm. 344 Journ. Hattori Bot. Lab. No. 68 I 990

gametophytic characters. As far as we could examine, all Japanese specimens so far determined as "D. jlexicaule" by previous authors belong to D. crispatissmum. D. jlexicaule can be distinguished from D. crispatissimum by the usually not robust plant size, leaves with sheathing base rather suddenly narrowed into short subula, cells of dorsal surface of costa usually longer than the adjacent lamina cells. Detailed discussion see Frisvoll (1985).

7. Ditrichum sekii Ando et Deguchi ex Matsui et Iwats., sp. novo (Fig. 14, Map 2b) Ditrichum sekii Ando et Deguchi in Ando et aI., Land and Life in Itsukushima: 340 (1975), nom. nudo In habitu Ditrichio ambiguumo Best. similis, sed marginibus foliorum vegetativorum bistratosis, costis in sectione transversa roundis differt. Plants small, densely tufted, yellowish green to dark green. Stems ca. 7 mm high, simple, with central strand. Leaves 1.2- 2.2 mm long, erect and ftexuose, from oblong-ovate base gradually tapering to long subula with obtuse apex; margins plane throughout, clearly bistratose except basal part; costa broad, occupying most part of subula, in cross-section biconvex, with distinct stereid bands on both sides of guide cells; cells at upper part of sheathing base quadrate to irregularly quadrate, basal cells quadrate to short rectangular. Dioicous. Perichaetial leaves with long sheathing base. Capsules erect, cylindric, slightly curved and asymmetric, urn ca. 1.5 mm long, reddish brown, smooth or weakly plicate when dry; seta 1.2 cm long, yellowish brown; peristome teeth pale orange, divided into two divisions to the base, ca. 0.3 mm long, densely papillose throughout; exothecial cells rectangular to narrowly hexagonal; annulus of two cell rows; stomata at apophysis; operculum short to elogate conic. Calyptra cucullate. Spores papillose, 10-12 /lm in diam. Chromosome number unknown. Representative specimens examined: JAPAN . HONSHU . Hiroshima-ken, Isl. Miyajima, Seki 3102 :" holotype (HIRO; isotype NI CH) . KyUSHU. Kumamoto-ken, Hitoyoshi, Mayebara s.n. (Musci japonici 10: 499 as Trichodon tenuifolius); Miyazaki-ken, Mt. Osuzu, Kuwahara 242 (NICH) . Fertility: All six specimens examined are fertile. Ecological notes: Two specimens wi th ecological data were collected on soil. Distribution: Japan (Honshu and Kyushu). Ditrichum sekii resembles D. ambiguum Best, a North American species, in the leaf and capsule shapes, but differs from D. ambiguum in the costa anatomy. The costa of D. sekii is clearly biconvex in cross-section and has strongly developed stereid bands on both sides of the guide cells, whereas that of D. ambiguum is somewhat piano-convex and has poorly developed or sometimes undeveloped stereid bands on the adaxial side of the guide cells.

8. Ditrichum pallidum (Hedw.) Hampe (Fig. l5, Map Ic) Flora 50: 182 (1867). TrichoslOmum pa/lidum Hedw., Spec. Muse.: 108 (1801) - Leptotrichum pa/lidum (Hedw.) Hampe in C. MueH ., Syn. Muse. 1: 451 (1848). Ditrichum longipes Sak., Bot. Mag. Tokyo 52: 129 (1938). Ditrichum currilucki Grout, Moss Fl. N. Amer. 1 (4): 253 (1939). Plants small, densely tufted, yellowish green. Stems 5- 10 mm high, simple, with central strands. Leaves 1-4 mm long, erect and ftexuose, from ovate to oblong, gradually tapering to long channelled subula; margins plane throughout, unistratose below, bistratose in upper part above the shoulder; costa broad, occupying most of the base of subula, in cross-section stereid T. MATSUI & Z. !WATSUKI: A taxonomic revision of Ditrichaceae of Japan 345

e m u p t T 0 "L ~ rIll

T ~ g !a 1'" f-20--1 1G}®m b ~/l'r q s f-20-\ I 0 0 I a

I:;; 1

FIG. IS . Dilrichum pal/idum (Hedw.) Hampe. a, plant. b, leaf. c, cells at leaf base. d, cells at upper part of sheathing base. e, cells at leaf apex. f- k, cross-sections of leaf. I, cross-section of stem. m, perichaetialleaf. n, capsule. 0 , operculum. p, peristome teeth. q, annulus. r, exothecial cells. s, stoma. t, spore. u, calyptra. a, b, e-f, k- u drawn from Matsui 524 (KOCH); c, d, g-j from holotype of D. longipes (MAK). Scale bars in /lm . bands distinct on both sides of guide cells; cells at upper part of sheathing base long rectangular, basal cells short-rectangular to rectangular, sometimes forming hyaline zone. Autoicous. Perichaetial leaves shorter than stem leaves, without sheathing base. Capsules inclined, rarely 346 Journ. Hattori Bot. Lab. No. 68 I 990 erect, slightly asymmetric, urn 1- 2 mm long, reddish brown, furrowed when dry; seta 1--4 cm long or longer, usually f1exuose, bright yellow; peristome teeth yellowish orange, divided into two divisions to the base, ca. 0.5 mm long, densely papillose throughout; exothecial cells rectangular, 10--30 Ilm wide; annulus of two cell rows; stomata at apophysis; operculum long conic. Calyptra long cucullate. Spores coarsely and sparsely papillose, (l5-)17- 22Ilm in diam. Chromosome number: n= 13 (AI-Aish & Anderson 1961),26 (Bryan 1956). Representative specimens examined: JAPAN. HOHSHU. Niigata-ken, Isl. Sado, Ikegami 557 (TNS); Shizuoka-ken, Saruwatari, Hashimoto 8 (TNS) ; Mie-ken, Kawasaki-mura, Koizumi 70408 (TNS); Hyogo-ken. Nishinomiya-shi, Ma/sui 659 (KOCH); Hiroshima-ken, Mt. Kanmuri, Higuchi 2265 (HIRO); Yamaguchi-ken, Tokuyama-shi, Suzuoka 3103 (TNS). SHIKOKU. Kochi-ken, Mt. Miyaji, Takaki s.n. (KOCH). KYUSHU. Fukuoka-ken, Yame-mura, Nabeshima 579 (TNS); Kumamoto-ken, Hitoyoshi, Mayebara s.n. - holotype of D. longipes (MAK ; isotype NICH) . Fertility: Among 14 specimens examined, 13 are fertile. Ecological notes: All six specimens with ecological data were collected on soil. Distribution: Japan (Honshu, Shikoku and Kyushu), Europe and North America. Ditrichum pa/lidum is characterized by several sporophytic characters: (I) strumose capsules with longitudinal furrows when dry; (2) long and bright yellow seta extending to 4 cm; and (3) coarsely and sparsely papillose spores. D. pallidum is very closely related to D. rhynchostegium. The distinction will be made under D. rhynchostegium.

9. Ditrichum difficile (Duby) Fleisch. (Fig. 16, Map la) Musci FI. Buitenz. 1: 300 (1904). Trichostomum dijJicile Duby in Morilzi, Syst. Verz. Zoll. Pfl. : 134 (1846). Leptotrichum affine C. Muell., Bot. Zcit. 5: 825 (1847) - Ditrichum affine (c. Muell.) Hampe, Flora 50: 182 (1867). Leptotrichum boryanum C. Meull., Syn. Musc. 1: 452 (1848) - Ditrichum boryanum (c. MueH.) Hampc, Flora 50: 182 (1867). Leptotrichum capense C. Muell., Syn. Musc. 1: 453 (1848) - Ditrichum capense (c. MucH.) O . Kuntze, Rev. Gen. PI. 2: 835 (1891). Leptotrichumflexifolium Mitt., Kew J. Bot. 8: 257 (1856) - Ditrichumflexifolium (Mitt.) Hampe, Flora 50: 182 (1867). Leplotrichum macleanum Rehm. in Kindb., Enum. Bryin. Exot.: 91 (1859), nom. nudo Trichostomum laxifolium Hook. f. et Wils., FI. Nov. Zel. 2: 72 (1854) - Leptotrichum laxifolium (Hook. f. et Wils.) C. Muell., Gen. Musc. Fr.: 313 (1900) - Ditrichum laxifolium (Hook. f. et Wils.) Mitt., Trans. R. Soc. Victoria 19: 51 (1882). Leptotrichum muelleri Hampe, Linnaea 28: 206 (1856) - Ditrichum muelleri (Hampe) Hampe, Flora 50: 182 (1867). Trichostomum oldjieldii Mitt. in Hook. f. , FI. Tasm. 2: 177 (1859) - Leptotrichum o/djieldii (Mitt.) Mitt., J. Linn. Soc. Bot. 4: 67 (1859) - Ditrichum o/djieldii (Mitt.) Mitt. in F. MucH., Fragrn. Phytogr. Austr. suppl. 11: 109 (1881 ). Leptotrichum plicatum C. Muell., Syn. Musc. 1: 446 (1848) - Ditrichum plicatum (C. MueH.) Hampe, Nuov. Giron. Bot. Ital. 4: 273 (1872). Trichostomum setosum Hook. f. et Wils., FI. Nov. Zel. 2: 73 (1854) - Leptotrichum setosum (Hook. f. et Wils.) C. Muell., Gen. Musc. Fr.: 313 (1900) - Ditrichum setosum (Hook. f. et Wils.) Reichdt., Reise Oesterr. Freg. Novara Bot. 1: 172 (1870). Diaphanophyllum vallis-gratiae Lindb., Oefv. K . Vet. Ak. Foerh. 19: 605 (1868) - Leptotrichum valis-gratiae (Lindb.) Hampe in Jaeg., Ber. S. Gall. Naturw. Ges. 1871 - 72: 462 (1873) - Ditrichum vallis-gratiae (Lindb.) Hampe, Flora 50: 182 (1867). T. MATSUI & Z. IWATSuKI: A taxonomic revision of Ditrichaceae of Japan 347

FIG. 16. Ditrichum difficile (Duby) Fleisch. a, plant. b-c, leaves, d, cells at upper part of sheathing base. e, leaf apex. f- i, cross-sections of leaf. j, cross-section of stem. k, capsule. I, operculum. m, peristome tooth. n, exothecial cells. 0, stoma. p, caly­ ptra. a- i and k drawn from holotyp of D. jormosicus (NICH); j, I- p from lwatsuki et al. 2594 (NICH). Scale bars in j.lm.

Leptotrichum capense C. Muell. var. vallis-gratiae C. Muell. in Par., Ind. Bryol.: 392 (1896), nom. nudo DUrichum capense (c. Muell.) Par. var. vallis-gratiae Par., Ind. Bryol.: 392 (1896), nom. nudo Leptotrichum semi-lunare C. Muell., Hedwigia 37: 112 (1898) - Ditrichum semi-lunare (c. Muel!.) Par., (nd. Bryol. suppl.: 132 (1900). Leptotrichum viride C. Muell., Hedwigia 37: 113 (1898) - DUrichum viride (c. MueH.) Par., Ind. Bryol. supp!.: 132 (1900). Ditrichum baileyi C. Muell. in Watts et Whitelegge, Proc. Linn. Soc. N. S. Wales, suppl. 27: 35 (1902), nom. nudo Ditrichumflavipes C. Muell. ex Rodway, Pap. Proc. R. Soc. Tasmania 19: 92 (1913). Ditrichum jormosicum Nog., J. Jap. Bot. 14: 397 (1938). Plants small, loosely to densely tufted, yellowish green to dark green. Stems 5- 20 mm high, simple or rarely branched, with central strands. Leaves 1--4 mm long, occasionally becoming 348 Journ. Hattori Bot. Lab. No. 68 I 990

• DiuichuII heteromallu• • Ditrichum sekii ry' i' ... 't-·----i-----:.--~ ' ~Ji___;_- . /

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• Dj ir ichum macrorhynchum ... Saelania glaucescens /

MAP 2 T. MATSUI & Z. IWATSUKJ: A taxonomic revision of Ditrichaceae of Japan 349 longer, erect and flexuose, ovate to oblong, suddenly narrowed to long channelled subula; margins plane throughout, unistratose below, bistratose above shoulder; costa broad, occupying about 1/2 of the base of subula and most part of upper subula, with distinct stereid bands on both sides of guide cell s; cells at upper part of sheathing base quadrate to rectangular and cells at basal part of sheathing base rectangular except several marginal rows where narrowly rectangular, forming more or less hyaline zone. Autoicous. Perigonia formed a short distance below perichaetia; perichaetialleaves with a long sheathing base, 4-5 mm long; cells from upper part of sheathing base short to elongate rhomboidal. Capsules erect, cylindric, slightly curved, asymmetric, occasionally symmetric in shorter capsules, urn 1-3 mm long, reddish brown, becoming appressed and flattened when dry and empty, rarely weakly furrowed when dry; seta l-4cm long, yellowish brown to reddish brown, rarely bright yellow; peristome teeth yellowish orange to pale orange, divided into two divisions to the base, 0.5-1.0 mm long, densely papillose throughout; exothecial cells rectangular; annulus of two cell rows; stomata at apophysis; operculum short to high conic. Calyptra cucull ate. Spores finely papillose, 12- 17/lm in diam. Chromosome number: n = 13 (Ramsay 1974), 13 + m (Ramsay 1974). Illustrations: Noguchi 1938: 398, Fig. 6 (as D. formosicum); Seppelt 1982b: 155, Figs. 1- 10. Representative specimens examined: TAIWAN. Mt. Li, Wang 1907 (NICH); Mt. Taihei, Noguchi 6647 - holotype of D. formosicum (NICH); Mt. Ali, Iwatsuki & Sharp 1378 (NICH). Fertility: All five specimens examined are fertile. Ecological notes: Among two specimens with ecological data, one was collected on soil , and one on cliff. Distribution: Taiwan, Soviet Asia, Indonesia, India, M adagascar, South Africa, South America, Australia and New Zealand. Ditrichum difficile is characterized by (I) long yellowish seta, (2) asymmetric capsules, (3) long peristome teeth and (4) fi nely papillose spores. Ditrichum difficile is quite similar to D. rhynchostegium. The distinction will be made under D. rhynchostegium.

10 . Ditrichum rhynchostegium Kindb. (Fig. 17, Map 2e) Rev. Bryo!. 37: 14 (19 10). Ditrichum henryi Crum et Anders., J. Elisha M itchell Sci. Soc. 72: 289 (1956). Plants small, loosely to densely tufted, yellowish green to dark green. Stems 5- 20 mm high, simple or rarely branched, with central strands. Leaves 1-4mm long, occasionally becoming longer, erect and flexuose, ovate to oblong, gradually tapering to long channelled subula; margins plane throughout, unistratose below, bistratose above shoulder; costa broad, occupying about 1/2 of the base of subula and most part of upper subula, in cross-section stereid bands distinct on both sides of guide cells; cells at upper part of sheathing base quadrate to rectangular, cells at leaf base rectangular except several marginal rows where narrowly rectangular, sometimes forming hyaline zone. Autoicous. Perichaetialleaves with a long, sheathing base, 4-5 mm long; cells from upper part of sheathing base short to elongated rhomboidal. Capsules erect, cylindric, slightly curved and asymmetric, occasionally symmetric in shorter capsules, urn 1- 3 mm long, reddish brown, becoming appressed and flattened when dry and empty, rarely weakly furrowed when dry; setae 1-4 cm long, reddish brown, rarely bright yellow to yellowish brown; peristome teeth yellowish orange to pale orange, divided into two divisionS to the base, 0.5- 1.0mm long, densely papillose throughout; exothecial cells rectangular; annulus of two cell-rows; stomata at apophysis; operculum high conic. Calyptra long cucullate. Spores fine ly, vermiculately and striately papillose, 12- 17 /lm in di am. Chromosome number: n = 13 (Crum & Anderson 1956). 350 Journ. Hattori Bot. Lab. No. 68 I 9 9 0

g a o~. u T ..• Io ••.:. .

t

f-20-1

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To ~ T 1 0 .L'" d~ FIG. 17. Ditrichum rhynchostegium Kindb. a, plant. b-<:, leaves. d, cells at leaf base. e, cells at upper part of sheathing base. f, cells at middle part of subula. g, cells at leaf apex. h- k, cross-sections of leaf. I, cross-section of stem. m, perichaetial leaf. n, capsule . 0 , operculum. p, peristome teeth. q, annulus. r, exothecial cells. s, stoma. t, spore. u, calyptra. a, b, d- k, n drawn from to po type of D. rhynchostegium (us); c, I, m, o-u from Deguchi 16630 (KOCH). Scale bars in !lm. T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 351

FIG. 18. Spores. a, Ditrichum difficile (lwatsulci et aI., NICH). b, D. pallidum (Matsui 524, KOCH). c, D. rhynchostegium (holotype of D. henryi, DUKE). Scale bar: 5/lm.

Representative specimens examined: JAPAN. HOKKAlDO . Mt. Apoi, Furuki 5463 (IURO). HONSHU . Aomori-ken, Mts. Hakkohda, Deguchi 16299 (KOCH); Akita-ken, Mt. Kurikama, Kanno 266 (NICH); Miyagi-ken, Sendai-shi, Noguchi 4557 (N ICH); Niigata-ken, Kanose, lkegami 78188 (NI CH); Toyama-ken, Mt. Kurobe, Sasaoka s.n. (NICH); lshikawa-ken, Mt. Hakusan, Ando 26227 (H1RO); Saitama-ken, Mt. Ryohgami, Shimizu s.n. (NICH); Ibaraki-ken, Mt. Hanazono, Watanabe 23374 (NICH); Aichi-ken, Mt. Sanage, Amano 8786 (NICH); Mie-ken, Mt. Gozaisho, Iwalsuki & Zanten 20 (NICH); Shiga-ken, Mt. Keikan, Deguchi 23975 (KOCH); Wakayama-ken, Mt. Ryuumon, Deguchi 8961 (KOCH); Osaka-fu, Katsuo-dera, Mizutani 2455 (NI CH); Okayama-ken, Niimi-shi, Higuchi 6157 (HIRO); Tottori-ken, Mt. Daisen, Mizutani s.n. (NICH); Hiroshima-ken, Isl. Miyajima, Matsui 517 (KOCH). SHIKOKU . Ehime-ken, Shionashi, Deguchi 23006 (KOCH); Tokushima-ken, Mt. Tsurugi, Deguchi 21123 (KOCH); Kochi-ken, Mt. Shiraga, Deguchi 20533 (KOCH). KV USHU. Nagasaki-ken, Is!. Fukue, Iwatsuki 6867 (NIC H); Kumamoto-ken, Aida, Mayebara 1010 (NICH); Miyazaki-ken, Mt. Oyaji, Iwatsuki & Minamidani 12362 (NICH); Kagoshima-ken, Mt. Shibi, Mizutani s.n. (NICH). KOREA . Is!. Quelpart, Choe 123 (NICH). TAIWAN. Mt. Ali , Noguchi 6695 (NICH) . Fertility: Among 325 specimens examined, 319 are fertile. Ecological notes: Among 173 specimens with ecological data, 157 were collected on soil, IS on rocks (inc!. two on stone walls and one on limestone), and one on rotten stump. Distribution: Japan (Hokkaido, Honshu, Shikoku, Kyushu and Ryukyu), Taiwan, Korea and North America. Although no type specimen of Ditrichum rhynchostegium was available for the present study, we were able to examine the holotype of D. henryi and the topotype specimen of D. rhychostegium cited in Robinson (1966) who first discussed the identity of D. rhynchostegium and D. henryi. Dr. H. Crum, one of the authors of D. henryi, later checked the type spcimen of D. rhynchostegium and confirmed Robinson's treatment (pers. corn. Or. H. Robinson to Or. H. Oeguchi) and accepted it in his work (Crum & Anderson 1981). A great majority of Japanese specimens so far recorded under the name " D. pallidum" belong to D. rhynchostegium Kindb. which is closely related to D. pallidum. The plants of the two species are occasionally intermixed with each other in lowland of Japan. D. rhynchostegium is also quite similar to D. difficile. The distinguishing characters of the above three species are shown below. 352 Journ. Hattori Bot. Lab. No. 68 I 990

D. dijJicile D. pal/idum D. rhynchostegium

Costa gradually narrow suddenly wide gradually narrow from base at shoulder from base Perichaetial leaf with sheathing with non­ with sheathing base sheathing base base Color of seta bright yellow bright yellow reddish brown (rarely bright yellow to yellowish brown) Surface of spore finely papillose coarsely and vermiculately and (Fig. 18a) sparsely papillose striately papillose (Fig. 18b) (Fig . 18c) Spore size 12- 17 Jlm 12- 17 Jlm 16-25Jlm

D. rhynchostegium is the commonnest species among the Japanese species of Ditrichum.

11 . Ditrichum divaricatum Mitt. (Fig. 19, Map 2d) Trans. Linn. Soc. Bot. ser. 2, 3: 155 (189 1). Ditrichum divaricatum Mitt. var. exaltatum Card., Bull . Herb. Boiss. ser. 2, 8: 155 (1908), syn. novo Plants large as the genus, densely tufted, brownish green to dun green. Stems 10-40 mm high, simple or sometimes branched, with central strands. Leaves 4-6mm long, flexuose, spreading, occasionally more or less erect, long sheathing, abruptly contracted to base of subula; margins plane throughout, unistratose below, bistratose above the shoulder; costa broad, occupying most of mamillose subu1a, in cross-section with distinct stereid bands on both sides of guide cells; cells of upper part of sheathing base irregularly quadrate, thick-walled, those of leaf base elongate to linear-rectangular, thin-walled. Filamentous gemmae present in leafaxils. Autoicous. Perichaetial leaves with a long, sheathing base. Capsules erect, cylindric, symmetric, rarely slightly asymmetric, urn 2- 3 mm long, brown; seta 1- 3 cm long, yellow to yellowish brown, straight to slightly flexuose; peristome teeth light brown at base, whitish above, divided into two divisions to the base, sometimes imperfectly split, ca. 0.4 mm long, papillose, with ornamentation obliquely striolate; exothecial cells rectangular; annulus of two cell rows; stomata at apophysis; operculum high conic. Calyptra long cucullate. Spores slightly papillose, 13- 16 ~m in diam. Chromosome number unknown. lllustrations: Takaki 1957: 96, Fig. I; Noguchi 1987: 113, Fig. 42. Representative specimens examined: JAPAN. HONSHU . Aomori-ken, Mts. Hakkohda, Deguchi 16106 (KOCH); Saitama-ken, Mts. Chichibu, Shimizu s.n. (NICH); Nagano-ken, Mt. Yatsuga-take, Furuki 1971 (HIRO); Yamanashi-ken, Mt. Fuji, Deguchi 28068 (KOCH); Shizuoka-ken, Mts. Akaishi, Oizuru 525 (NICH); Tochigi-ken, Yumoto, Shirane-san, Bisset s.n. - lectotype of D. divaricatum (NY, selected here); Gifu-ken, Maeda 41 (NICH); Nara-ken, Mt. Ohmine, Nakajima 7542 (NICH) . SHIKOKU. Ehime-ken, Mt. Ishizuchi, Oti 13935 (NICH): KYUSHU. Ohita-ken, Mt. Sobo, Kuwahara 1204 (NICH). KOREA. Quelpaert, Hallaisan, Faurie 606-isotype of D. divaricatum var. exaltatum (KYO) . Fertility: Among 72 specimens examined, 54 are fertile. Ecological notes: Among 30 specimens with ecological data, 16 were collected on soil, and 14 on rocks. Distribution: Japan (Hokkaido, Honshu, Shikoku and Kyushu) and Korea. Ditrichum divaricatum was described by Mitten (1891) based on two specimens: T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 353 s r n 0·::'... . 1-20-1

m~.', Io 1'" 1--5 0 O--{

FIG. 19. Ditrichum divaricatum Mitt. a, plant. b, leaf. c, cells at middle part of sheathing base. d, cells at upper part ' of sheathing base. e, cells at leaf apex. f- i, cross-sections of leaf. j, cross-section of stem. k, axilally gemma. I, capsule. m, operculum. n, peristome teeth. 0, annulus. p, exothecial cells. q, stoma. r, spore. s, calyptra. a--<;, g,j- s drawn from Deguchi 22770 (KOCH); d- f, h- i from lectotype of D. divaricatum (NY). Scale bars in jlm. 354 Journ. Hattori Bot. Lab. No. 68 1 990

" Shiranesan, with young seta in August; Nantaizan, Septermber, with old capsules, Bisset." A specimen labeled "Shiranesan, Yumoto, 8/87, Bisset" is selected here as the lectotype, since it contains Mitten's pencil drawings, and many sporophytes, which agrees with the description, than another specimen. Ditrichum divaricatum has distinct gametophytic characters such as the leaves with long sheathing base, and the distinctly mamillose subula. In addition to those characters, Oeguchi (1981) reported filamentous gemmae, which have never been found in Ditrichum. Ditrichum divaricatum is closely related to D. cylindricarpum (c. Muell.) F. Muell. of Australia, New Zealand and South America. D. divaricatum has yellow to yellowish brown setae and filamentous gemmae in leafaxils, whereas D. cylindricarpum has reddish-brown setae and no filamentous gemmae. However we feel that these distinguishing characters are not always reliable. Further study with more specimens of both species is needed.

5. Trichodon Schimp. Coroll.: 36 (1856). Type species: Trichoslomum cylindricum Hedw. [= Trichodon cylindricus (Hedw.) Schimp.]. Trichodon, a monotypic genus, is closely related to Ditrichum and many authors (Grout 1936, Noguchi 1987, et al.) placed it in the genus Ditrichum. A taxonomic study of the genus Trichodon in North America was made by Ireland (1978), but he did not discuss any sporophytic characters to distinguish Trichodon and Ditrichum. Trichodon differs from Ditrichum by its squarrose leaves and incurved peristome teeth.

I. Trichodon cylindricus (Hedw.) Schimp. (Fig. 20, Map Id) CoroB.: 36 (1856). Trichostomum cylindricum Hedw., Spec. Musc.: 107 (1801) - Dicranum cylindricum (Hedw.) Srn., FI. Brit. 3: 1221 (1804) - Didymodon cylindricus (Hedw.) Wahlenb., Ft. Suec. 2: 754 (1826) - Ceratodon cylindricus (Hedw). Fuernr., Flora 12: 31 (1829) - Aongstroemia cylindrica (Hedw.) C. Muel!. , Syn. Muse. I: 441 (1848) - Leptolrichum cylindricum (Hedw.) Vent et Bott., Atti Soc. Critt. Ita!. 2, 3: 198 (1884) - Ditrichum cylindricum (Hedw.) Grout, Moss FI. N. Am. I: 48 (1936) . Trichodon lenuifolius Lindb., Oefv. K. Sv. Vet. Akad. Foerh. 21 : 225 (I864). Plants small, in loosely caespitose tufts. Stems to 20 mm high, mostly simple, with central strands. Leaves 1- 2 mm long, oblong to oblong-ovate with sheathing base, abruptly narrowed to a long, linear, roughened, wide-spreading subula; margins plane and unistratose at base, bistratose above, crenate-serrulate at the subula, rough at back, in cross-section stereid bands weakly differentiated at abaxial side of guide cells; cells of the leaf base oblong, lax, thin-walled, smooth, 20-43 /.lm long, those at the upper part of sheathing base short-rectangular to rectangular, 12- 33/.lm long. Rhizoidal tubers present. Dioicous. Capsules erect to suberect, narrowly cylindric, usually curved, asymmetrical, 1 mm long, smooth when dry and empty; seta ca. 10 mm Icing, slender and flexuose; peristome teeth with a low basal membrane, the 16 teeth incurved and not twisted, ca. 0.3 mm long, spilit nearly to the base into 2 filiform segments, finelly papillose; operculum high conic, ca. 0.4 mm long. Calyptra cucullate. Spores somewhat papillose, 14-16/.lm in diam. Chromosome number: n= 13 (Steere 1954). Illustrations: Deguchi 1982: 10, Fig. 2; Noguehi 198 7: 119, Fig. 458 (as Dilrichum cylindricum). Representative specimens examined: JAPAN . HONSHU. Saitama-ken, Mt. Mae-shiroiwa, Noguchi T. MATSUI & Z. !WATSUKI : A taxonomic revision of Ditrichaceae of Japan 355

t-20-1 b

FIG. 20. Tirchodon cy/indricus (Hedw.) Schimp. a, plant. b-f. leaves. g, cells at leaf base. h, cells at upper part of sheathing base. i, cells at leaf apex. j-o, cross-sections of leaf. p, cross-section of stem. q, perichaetial leaf. r- s, capsules. t, peristome tooth. u, exothecial cells. v, stoma. w, spores. x, rhizoidal tuber. a-q and s drawn from Deguchi 21093; r, t- x drawn from Noguchi 35284 (NICH) . Scale bars in }lm.

35284 (NICH). SHIKOKU. Tokushima-ken, Mt. Tsurugi, Deguchi 24110 (KOCH). Fetrility: Among six specimens examined, three are fertile. Ecological notes: Among five specimens with ecological data, three were collected on soil , and two on rocks (limestone). Distribution: Japan (Honshu, Shikoku), central Asia, Europe and North America. Trichodon cylindricus is characterized by the squarrose leaves, weakly developed stereid bands, asymmetric capsules and incurved peristome teeth.

6. Pleuridium Rabenh., nom. cons. Deutschl. Krypt. F1. 2: 79 (1848); non Pleuridium Brid. (1819). Type species: Phascum subulatum Hedw. [ = Pleuridium subulatum (Hedw.) Rabenh.]. Plants in loose tufts. Stems short, 3- 7 mm high, occasionally branched, epidermal layer 356 Journ. Hattori Bot. Lab. No. 68 I 990 differentiated, central strands present. Leaves appressed to erect-spreading, from ovate to oblong-ovate, tapering gradually or rapidly to subula, weakely channe1ed; margins plane, somewhat incurved above, smooth to serrulate at the apex; costa occupying most part of the subula, stereid bands differentiated on adaxial and abaxial sides of guide cells; cells of sheathing base oblong-rhomboidal to rectangular at middle part, rectangular at the base. Autoicous. Capsules cleistocarpus, globose with apiculus; setae short. Calyptra cucullate. Spores finely papillose. According to Snider and Margadant (1973), Pleuridium Brid. includes species that have long been placed in Archidium. It is highly desirable to conserve Pleuridium Rabenh. against a previous homonym, Pleuridum Brid. in order to validate current usage.

KEY TO THE SPECIES 1. Leaves appressed, 0.5-0.9 mm long, from ovate base abruptly elongate to short subula ...... I . P. julaceum (p. 356) I. Leaves erect to erect spreading, 1- 3 mm long, from oblong-ovate base gradually to abruptly subulate ...... 2. P. subulatum (p. 357)

1. Pleuridiumjulaceum Besch. (Fig. 21, Map Id) J. de Bot. 12: 294 (1898). Stems 4-5 mm high. Leave 0.5-0.9 mm long, appressed to the stems, from ovate base abruptly elongate to short subula, weakly channeled; margins plane; costa excurrent, filling most of the subula, stereid bands differentiated on abaxial side of guide cells; cells at upper

F IG. 21. Pleuridiumjulaceum Besch. a- b, plants. c-e, leaves. f, cells at leaf base. g, cells at upper part of sheathing base. h, cells at leaf apex. i- k, cross-sections of leaf. Drawn from isotype of P. julaceum (NICH). Scale bars in Ilm. T. MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 357 part of sheathing base quadrate to short rectangular, cells at leaf base rectangular. Autoicous? Seta 0.5 mm long. Capsules spherical?, I mm long. Calyptra cucullate, 0.8 mm long. Chromo­ some number unknown. Illustration: Noguchi 1987: lOS, Fig. 40B. Specimen examined: JAPAN. HONSHU. Tokyo-to, Koishikawa Bot. Garden, Malsumura? s.n. - isotype of P. jlllacellm (NICH). Fertility: Isotype specimen examined is fertile. Ecological notes: Ecological data unknown. Distibution: Japan (Honshu), endemic. Pleuridium julaceum was collected from Koshikawa Botanical Gerden of the University of Tokyo. Only one flattened c1eistocarpous capsule was found in the type specimen of P. julaceum. However, we could not examine detailed structures of the capsule. Acording to Ihsiba (1929), P. julaceum was found in Taiwan. We found only one specimen determined as "P. julaceum" from Taiwan (Taipei, Oct. 6, 1907, Tsunoda S. n., TNS), but this specimen belong to Brachymenium excite.

2. Pleuridium subulatum (Hedw.) Rabenh. (Fig. 22, Map Id) Deutsch!. Krypt. Fl. 2: 79 (1848). Phascum subulalum Hedw., Spec. Muse.: 19 (1801) - ASlOmum subulatum (Hedw.) Hampe, Flora 20: 285 (1837). Pleuridium acuminatum Lindb., Oefv. K. Sv. Vet. Akad. Foerh. 20: 406 (1863). Plants in loose, yellow green or yellow brown tufts. Stems 3- 7 mm high, occasionally branched, with central strands. Leaves 1- 3 mm long, loosely erect or erect-spreading, oblong-ovate, tapering gradually to abruptly subulate, weakly channeled; margins plane, somewhat incurved above, serrulate at the apex; costa occupying most part of subula, stereid bands clearly differentiated on adaxial and abaxial sides of guide cells; cells of sheathing base linear above, oblong-rhomboidal to rectangular at the shoulder, rectangular at the base. Paroicous or autoicous. Antheridia few, naked in axils of the upper leaves or in small bracts. Capsules globose with apiculus, 0.5- 1 mm in diameter; setae short, ca. 0.5mm long. Calyptra cucuUate, small, ca. 0.5 mm long. Spores finely papillose, 20-30 J.lm in diam. Chromosome number: n= 13 (Bryan 1956). Illustration: Noguchi 1987: lOS, Fig. 40C. Representative specimens examined: JAPAN. HONSHU . Niigata-ken, Is!. Sado, Homma 23433 (NICH); Fukushima-ken, Higuchi 9316 (NICH); Ibaraki-ken, Tomobe-cho, Izawa s.n. (NICH); Shiga-ken, Rittoh-cho, Deguchi 23999 (KOCH); Kyoto-fu, Mt. Hiei, Toyama s.n. (NICH); Mie-ken, Nabari-shi, Tailoh 63801 (NICH); Osaka-fu, Kaizuka-shi, Nakajima 2341 (NICH); Nara-ken, Nara-shi, Deguchi 5592 (KOCH); Hyogo-ken, Yatutomi-mura, Tetebe 560 (NICH); Okayama-ken, Mt. Takatsuma, Kiguchi 1829 (NICH); Hiroshima-ken, Yahata-shrine, Deguchi 14963 (KOCH); Yamaguchi-ken, Itone-mura, Matsumoto 485 (NICH). SHIKOKU. Ehime-ken, Sakura-machi, Oli 4468 (NICH); Kochi-ken, Mt. Kajigamori, Hidaka 176 (KOCH). KVUSHU. Fukuoka-ken, Mt. Kohra, Shikata s.n. (NICH). Nagasaki-ken, Is!. Hirado, Hiramalsu 46 (NICH); Kumamoto-ken, Hitoyoshi-shi, Mayebara s.n. (Musei Japonici Exisecati ser. 10, no. 485, HIRO). Fertility: All 27 specimens examined are fertile. Ecological notes: Among 17 specimens with ecological data, 16 were collected on soil, and one was collected on rock. Distribution: Japan (Honshu, Sh'ikoku and Kyushu), eastern Asia, Europe, North America. Pleuridium subulatum is characterized by the small plants, globose capsules and long subulate leaves. 358 Journ. Hattori Bot. Lab. No. 68 I 990

h

T o r\ :;:

b d

To ~ 1

FIG. 22. Pleuridium subulalum (Hedw.) Rebenh. a, plant. b-f, leaves. g, cells at sheathing base. h, leaf apex. i- m, cross-sections of leaf. n, cross-section of stem. 0, capsule. p, exothecial cells. q, stoma. r, spore. s, calyptra. Drawn from Deguchi 14963 (KOCH). Scale bars in JJ.m.

7. Pseudephemerum (Lindb.) Hag. K. Norsk. Vid. Selsk. Skrift. 1910: 45 (1910). Type species: Phascum axil/are Srn. [=Pseudephemerum nitidum (Hedw.) Reim.]. Pleuridium sect. Pseudephemerum Lindb., Oefv. K. Vet. Ak. Foerh. 21 : 583 (1865) - Pleuridium subgen. Pseudephemerum (Lindb.) Broth., Nat. PH . 1: 294 (1901). Pleuridiella ·Robins., J. Hattori Bot. Lab. 27: 125 (1964), syn. novo Pseudephemerum is characterized by lanceolate leaves, thin walled cells of sheathing base and c1eistocarpous capsules. Pseudephemerum is similar to Eccremidium subgen. Pseudopleuridium, but capsules of Pseudephemerum are not dehiscent near the equator. T. MATSUI & Z. !WATSUKI : A taxonomic revision of Ditrichaceae of Japan 359

1. Pseudephemerum nitidum (Hedw.) Reim. (Fig. 23, Map 2a) Verh. Bot. Ver. Brandenburg 74: 152 (1933). Phascum ni/idum Hedw., Spec. Musc.: 19 (1801) - Pleuridium nitidum (Hedw.) Rabenh., Deutschl. Krypt. Ft. 2: 79 (1848). Phascum axil/are Dicks. ex Srn., Engl. Bot. 14: 1036 (1802) - Pleuridium axil/are (Srn.) Lindb., Oefv. K. Vet. Ak. Foerh. 20: 407 (1863) - Pseudephemerum axil/are (Srn.) Hag., K. Norsk. Vid. Selsk. Skrift.

FIG. 23. ·Pseudephemerum nitidum (Hedw.) Hag. a, plant. b-e, leaves. f, cells at leaf base. g, cells at upper part of leaf. h, cells at leaf apex. i- k, cross-sections of leaf. I, cross-section of stem. m, capsule. n, calyptra. 0, exothecial cells. p, stoma. q, spores. a, h, I and o-q drawn from lwatsuki, Bryophyta Exsiccata Fasc. 3, no. 129 (NIGH); b--g, i- k and rn- n from Deguchi 5637 (KOCH). Scale bars in /lm . 360 Journ. Hattori Bot. Lab. No. 68 1 990

1910: 45 (1910). Ephemerella caldensis C. Mucll., Bot. Zeit. 17: 197 (1859) - Pleuridium caldensis (c. Muel!.) Lindb., Oefv. K. Vet. Ak. Foerh. 21 : 584 (1865) - Pseudephemerum caldense (c. Muell.) Broth., Nat. Pfl. ed. 2, 10: 177 (1924). Sporledera laxifolia Ren. et Card., Act. Soc. Linn. Bordeaux 53: 18 (1898) - Pseudephemerum laxifolia (Ren. et Card.) Ther., Recueil Pub!. Soc. Havraise, Etud. Div. 1932: 139 (1932). Pleuidiella colei Robins., J. Hattori Bot. lab. 27: 125 (1964) - Pleuridium colei (Robins.) Deguchi et Matsui, J. Jap. Bot. 60: 330 (1985), syn. novo Plants small, in caespitose patches. Stems 2- 3 mm high, simple or occasionally branched, with central strands. Upper leaves 1.4-2.3 mm long, 0.3- 0.5 mm wide, lanceolate, keeled above, with recurved margins that are denticulate by the projection of upper corner of marginal cells; costa excurrent, in cross-section epidermal layers of large, thin walled cells; cells of sheathing base elongate rectangular, becoming longer and wider, with thinner walls toward leaf base. Synoicous. Capsules immersed in perichaetialleaves, globose, with apiculum, 0.4-0.5 mm long, without differentiation of operculum; exothecial cells round quadrate, with smooth, thin walls; stomata phaneroporous, in 2- 3 concentric rows on lower half of capsule, seta ranging 0.08-0.12 mm long. Calyptra small, cucullate, sometimes with eroded base. Spores finely papillose, 35-40 Ilm in diam. Chromosome number: n = 13 (Smith & Newton 1968). Illustrations: Iwatsuki 1980: 130, Fig. I; Deguchi & Matsui 1985: 328, Fig. 1 (as Pleuridium colei); Noguchi 1987: 143, Fig. 54B. Specimens examined: JAPAN . HONSHU. Hiroshima-ken, Suzugamine, Deguchi 5637 (KOCH). KYUSHU. Nagasaki-ken, Is!. Fukue, Iwatsuki s.n. (Bryophyta Exsiccata Fasc. 3, no. 129, HIRO, NICH). Fertility: Two specimens examined are fe rtile. Ecological notes: Two specimens examined were collected on soil in rice fields. Distribution: Japan (Honshu and Kyushu), Assam, East Nepal, Europe and North America. Robinson (1964) described the genus Pleuridiella with a species P. colei, based on the lanceolate leaves and globose capsules with numerous stomata. Recently Deguchi and Matsui (1985) considered Pleuridiella as a synonym of Pleuridium, but the distinguishing characters of P. colei are quite similar to those of Pseudephemerum nitidum. Pleuridium colei should be reduced to synonymy of Pseudephemerum nitidum.

8. Eccremidium Wils. London J. Bot. 5: 450 (1846). Plants very small , pale to yellowish green, scattered or gregarious on bare soi l. Stems 1- 3 mm high, simple, with central strands. Leaves laxly ftexuose, concave, broadly ovate, and slenderly lance-acuminate; margins plane, entire or crenate-serrulate to serrate above; costa sometimes weak or absent at base, in cross-section stereid bands absent; cells of sheathing base thin-walled, hexagonal to oblong-rhomboidal above; comalleaves not much differentiated, erect, ftexuose or subsecund, elongate and subulate-acuminate from a broader base. Autoicous. Perichaetial leaves scarcely differentiated. Capsules c1eistocarpous, globose, dehiscent along equator line, marked by 2 rows of smaller, oblate cells; setae short; exothecial cells hexagonal, thin-walled. Calyptra small, conic-mitrate, roughened by projecting cell ends. Spores reticulate-papillose. Eccremidium is characterized by globose capsules with dehiscence line near the equator, costa without stereid bands and thin walled cells of sheathing base. Eccremidium is similar to Pleuridium and Pseudephemerum, but the dehiscence line of the capsule is significantly different. T . MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 361

KEY TO THE SPECIES 1. Leaves keeled, costa in 3-4 cell layers...... 1. E. brisbanicum (p. 361) I. L~aves plane, costa in 2 cell layers ...... 2. E. minutum (p. 362)

1. Eccremidium brisbanicum (Broth.) Stone et Scott (Fig. 24, Map 2a) J. Bryol. 7: 603 (1973). Archirium brisbanicum Broth., Oefv. Finsk. Vet. Soc. Foerh. 35: 35 (1893) - Nanomitrium brisbanicum (Broth.) Broth., Nat. Pfl. 10: 320 (1924) - Micromilrium brisbanicum (Broth.) Crosby, Bryologist 71 : liS (1968).

o f- 20--; {11

f-20--;

FIG. 24. Eccremidium brisbanicum (Broth.) Stone et Scott. a, plant. b-e, leaves. f, cells at leaf base. g, cells at upper part of sheathing base. h, cells at leaf apex. i- k, cross-sections ofleaf. I, cross-section of stem. m, capsule. n, exothecial cells. 0, stoma. p, spore. q, calyptra. a-{:, i- m and q drawn from Yamaguchi & Seki 4671 (HIRO); f- h, n- p from Deguchi 8808 (KOCH) . Scale bars in /lm. 362 Journ. Hattori Bot. Lab. No. 68 I 990

Pleuridium austro-subulatum Broth. ex Roth, Hedwigia 54: 269 (1924). Plants yellowish, loosely foliate. Stems 0.5-2 mm high, with thin-walled central strands. Leaves small below, longer above, long-acuminate from a concave, lanceolate base; margins entire at base, serrulate at middle to apex; cells of sheathing base rectangular to long rectangular, sometimes linear, 50-75(- 100)Jlm at basal part; costa indistinct at base, distinct at middle to apex, three to four cell layers in thickness, and round to ellipsoid in cross-section, stereid bands absent. Synoicous. Capsules globose, with equatorial dehiscence lines of narrow, elongated, thin-walled cells, stomata confined to lower half. Calyptra conic or wide bell shape and lobed at base, roughened by projecting cell ends, remains of archegonia often persisting at upper part. Spores 20-30 per capsule, reticulate papillose, 90-100 Jlm in diam. Chromosome number unknown. Illustration: Deguchi & Matsui 1986: 92, Fig. I. Representative specimens examined: JAPAN. HONSHU . Wakayama-ken, Susami-cho, Deguchi 8808 (KOCH, MELU). RYUKYU. Isl. Yonaguni, Yamaguchi & Seki 4671 (HIRO). Fertility: Three specimens examined are fertile. Ecological notes: Three specimens with ecological data were collected on soil (incl. two on moist soil). Distribution: Japan (Honshu and Ryukyu), Australia. The distinction between Eccremidium brisbanicum and E. minutum is discussed under the latter species.

2. Eccremidium minutum (Mitt.) Stone et Scott (Fig. 25, Map 2a) J. Bryol. 7: 603 (1973). Bruchia minuta Mitt. in Hook., Fl. Tasman. 2: 165 (1859). Bruchia whiteleggei C. Muell., Flora 71 : 10 (1888) - Sporledera whiteleggei (c. Muell.) Kindb., Enum. Bryin. Exot.: 95 (1889) - Ephemerum whiteleggei (C. MueH.) Broth. et Geh., Oefv. Finsk. Vet. Soc. Foerh. 37: 156 (1895). Plants yellowish, loosely foliate. Stems less than I mm high, with tbin-walled central strands. Leaves small below, longer above, long-acuminate from a concave, lanceolate base; margins entire at base, serrulate at apex; cells of sheathing base rectangular to long rectangular, 5G-65 Jlm long at basal part; costa indistinct, two cell layers in thickness, stereid bands absent. Synoicous. Capsules globose, with equatorial dehiscence lines of narrow, elongate, thin-walled cells, stomata confined to the lower half. Calyptra conic or wide bell shape and lobed at the base, roughened by projecting cell ends, remains of archegonia often persisting at upper part. Spores reticulate pepillose, 65- 100(- 140) Jlm in diam. Chromosome number unknown. Illustrations: Iwatsuki & Takaki 1979: 99, Fig. 2; Noguchi 1987: 108, Fig. 41B. Specimens examined: JAPAN. HONSHU . Aichi-ken, Toyohashi-shi, Takaki s.n. (Bryophyta Exsiccata, Fasc. 4, no. 160, H1RO); Hiroshima-ken, Isl. Miyajima, Seki s.n. (HIRO). Fertility: Two specimens examined are fertile. Ecological notes: Two specimens with ecological data were collected on wet soil. Distribution: Japan (Honshu) and Australia. Eccremidium minutum is closely relataed to E. brisbanicum in many features, but E. brisbanicum has keeled leaves with thick costa of 3-4 cell-layers which is rounded to ellipsoid. in cross-section. Scott and Stone (1976) mentioned that E. brisbanicum differs from E. minutum in the long stem (2- 3 mm) and in spores measuring 100-140 jJ.m . But the Japanese materials of these species are similar to each other by the spore size, being 90-100 jJ.m in E. brisbanicum, and to 65- 100 jJ.m in E. minutum. T . MATSUI & Z. IWATSUKI: A taxonomic revision of Ditrichaceae of Japan 363

FIG. 25. Eccremidium minutum (Mitt.) Stone et Scott. a, plant. b-d, leaves. e, cells at leaf base. f, cells at upper part of sheathing base. g, cells at leaf apex. h- j, cross-sections of leaf. k, cross-section of stem. I, capsule. m, exothecial cells. n, stoma. 0 , spore. p, calyptra. Drawn from Seki s.n. (HIRO). Scale bars in Jlm.

SPECIES NOT AVAILABLE for STUDY I. Ceratodon purpureus (Hedw.) Brid. var. formosicus Card., Beih. Bot. Centralbl. 19: 100 (1905). We could not located the type of this species. 2. Ditrichum astomoides Limpr., Laubm. Deutschl. 1: 511 (1887). Sakurai (1938) recorded this species from Japan. Noguchi (1987) cited that the specimens of this species are not located in Sakurai's collection in MAK. Also we were not able to find any 364 Journ. Hattori Bot. Lab. No. 68 I 990 specimen referable to D. astomoides in many Ditrichum specimens examined from Japan.

G ENERA AND SPECIES EXCLUDED FROM THE DITRICHACEAE I. Garckea C. Muell., Bot. Zeit. 3: 865 (1845). Garckea is characterized by the short seta, rostrate operculum, lanceolate peristome teeth that are fused at the apex, and lack of stomata. Fleischer (1904) and Brotherus (1924) placed Garckea in the Ditrichaceae. However, the characters mentioned above are quite different from those of other genera of the Ditrichaceae. Where as these characters are common to the Dicranaceae. Therefore, we prefer to place Garckea in the Dicranaceae. 2. C . MueH. , Bot. Centralbl. 7: 345 (188 1). Wilsoniella is characterized by its large, thin-walled cells, weakly differentiated stereid bands, costa ending below apex, rostrate operculum and spirally striolate peristome teeth. Wilsoniella was placed in the Dicranaceae by Fleischer (1904) and Brotherus (1924). Recently, it was transferred to the Ditrichaceae by Buck (1979), but gametophytic characters are similar to Dicranella or Microdus, which are members of the Dicranaceae. Moreover, surface structure of the peristome teeth of Wilsoniella is quite different from other genera of the Ditrichaceae. Therefore, we prefer to place Wilsoniella in the Dicranaceae. 3. Ditrichum mayebarae Sak., Bot. Mag. Tokyo 53: 62 (1939). After examination of the type material, we found that Ditrichum mayebarae has a similar habit to D. divaricalum, and similar leaves to D. heteromallum. However, D. mayebarae has Dicranum-type peristome teeth, and has capsules without stomata. These characters suggest that D. mayebarae is a member of the Dicranaceae, especially of Dicranella . Therefore, the following new combination is proposed:

Dicranella mayebarae (Sak.) Matsui et Iwats., comb. novo Ditrichum mayebarae Sak., Bot. Mag. Tokyo 53: 62 (1939). Type: Japan, Kumamoto-ken, Aida, Mayebara s.n., herb. Sakurai 9328 (holotype in MAK ; isotype in NI CH). 4. Ceratodon perplexans Card., Bull. Herb. Boiss. ser. 2, 7: 716 (1907). Our examination of the type specimen indicates that C. perptexans is characterized by its (I) broad-lanceolate to lanceolate leaves, (2) recurved leaf margins, (3) quadrate to rounded cells, and (4) blunt papillae on cells. These characters suggest that C. perplexans is a member of Didymodon in the Pottiaceae, and thus should be called Didymodon constrictus (Mitt.) Saito.

Didymodon constrictus (Mitt .) Saito, J. Hattori Bot. Lab. 39: 514 (1975). Ceratodon perp/exans Card., Bull. Herb. Boiss. SeT. 2, 7: 716 (1 907), syn. novo Type: Korea, Fusa n, Faurie 277 (isotype in KYO).

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