Hattoria 12: 51–92. 2021

Re-examination of the Symphyodon perrottetii () and related species in East and Southeast Asia

Hiroyuki AKIYAMA1, 2, Rimi REPIN3 & Monica SULEIMAN4

1 Museum of Nature and Human Activities, Hyogo, Yayoigaoka-6, Sanda, Hyogo 669–1546, Japan 2 Institute of Environmental Sciences, University of Hyogo, Yayoigaoka-6, Sanda, Hyogo 669–1546, Japan 3 Research and Education Division, Parks, 88806 , Sabah, 4 Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400 Kota Kinabalu, Sabah, Malaysia Author for correspondence: Hiroyuki AKIYAMA, [email protected]

Abstract Re-examination of morphological features and newly executed molecular analysis based on chloroplast (rbcL, rps4, and trnL-F) and nuclear (ITS1&2) genomes revealed unexpected diversity in Symphyodon perrottetii Mont. s. l. (Symphyodontaceae) in East and Southeast Asia. Japanese previously recognized as S. perrottetii are shown to include at least five different species: S. asper (Mitt.) A.Jaeger, S. copelandii Broth., S. gollanioides Nog., S. recurvomarginatus (Dixon & Sakurai) H.Akiyama, comb. nov. and S. rheophilus H.Akiyama, sp. nov. In addition, previous Japanese record of S. echinatus (Mitt.) A.Jaeger is actually S. asper. Symphyodon longispinosus H.Akiyama, sp. nov. from southwestern China is newly described. Subsequently, S. perrottetii s. str. is excluded from China and Japan and has its revised distribution range from southwest India (type locality of S. perrottetii), Sri Lanka, central , and . Symphyodon merrillii Broth., once regarded as a synonym of S. perrottetii, is again recognized as a distinct species with its distribution in Malesia and Indochina. Symphyodon ovalifolius H.Akiyama & M.Suleiman, sp. nov. is described and classified under the subgenus Parasymphyodon H.Akiyama & M.Suleiman, subg. nov. Rheoshevockia fontana Ignatov, W.Z.Ma & D.G.Long is again demonstrated to have a unique systematic position sister to the genus Symphyodon.

Introduction The bryophyte genus Symphyodon Mont. was established by Montagne (1841) with a single species, S. perrottetii Mont. from Neel-Gherries (＝Nilgiri Hills), southwestern India. The most important traits of S. perrottetii are cylindrical, erect, often distinctly echinate capsules, and reduced endostome. Müller (1850–1851) placed Symphyodon perrottetii under the genus Neckera allied with

51 N. angusta Müll.Hal., a species from Bombay, South India. Mitten (1859) treated Symphyodon as a section of the genus Stereodon (Brid.) Brid., and recombined the above two species and recognized a total of eight species in the section (including five new species based on specimens from Nepal, Khasia and Sikkim). Among them, S. planulus Mitt. was later designated as a member of the genus Aptychella (Broth.) Herzog (formerly as Clastobryopsis M.Fleisch.) as A. planula (Mitt.) M.Fleisch. Jaeger & Sauerbeck (1878) recognized the genus Symphyodon with eight species, mainly following Mitten’s (1859) sectional treatment of Symphyodon. Since then, the number of species in the genus has increased as results of several regional studies (Noguchi 1972; Tixier 1972; Gangulee 1976; Steere 1982; Buck & Ireland 1992; He & Snider 1992, 2000; Churchill 1994; Akiyama 2017a; He & Khang 2012). In a taxonomical revision of Symphyodon, He & Snider (2000) reduced the number of species from 27 known at that time to 15. This is currently considered to be the most reliable literature. After He & Snider’s (1992, 2000) revision, two additional species have been published: Symphyodon leiocarpus H.Akiyama & H.Tsubota (Akiyama & Tsubota 2009) from northern Thailand and S. chrysobasilaris (Broth.) W.Kim & H.Akiyama (Kim et al. 2020) from Hawaii. Thus, a total of 17 species are now recognized for the genus. However, Symphyodon is still one of the most difficult groups to define species delimitations because of little morphological difference among the species. The characters that are considered to be important in species identification are highly variable depending on growth conditions, and developmental stages, such as sizes, branching patterns, leaf shapes, and degree of laminal cell proration. Along with the genera Chaetomitriopsis M.Fleisch., Dozy & Molk., Dimorphocladon Dixon, Phyllodon Schimp., and Rheoshevockia Ignatov, W.Z.Ma & D.G.Long, Symphyodon has usually placed in the Symphyodontaceae M.Fleisch. (Goffinet et al. 2008; Ma et al. 2018; Kim et al. 2020). These genera are commonly characterized by a short, bifid costae, limited alar regions composed of small, quadrate to short-rectangular cells, and more or less prorate (rarely smooth) laminal cells. In nearly all of the genera in the family, morphological differentiation between stem leaves and branch leaves is highly correlated to branching patterns. The species with bi-pinnately to tri-pinnately branching patterns tend to have highly differentiated stem and branch leaves; in contrast, the species with irregularly and sparsely branching patterns have less differentiated leaves. In addition, some members of the Symphyodontaceae (Chaetomitrium and Symphyodon) are characterized by strong spiny protuberances or spines on the capsule walls and/or the upper part of the setae. Most of the species are epiphytes in slightly moist warm temperate to subtropical , but some are rheophytes in such genera as Phyllodon and Rheoshevockia. Sporophytes have not been found in Rheoshevockia, and morphology of capsules is remained ambiguous. Nevertheless, the latter genus is thought to be most closely related to Symphyodon (Ma et al. 2018). As mentioned above, 18 species of Symphyodon are now recognized in the world. Except for S. imbricatifolius, which is known only in Central and northern South America and S. pygmaeus (Broth.) S.He & Snider, a pan-tropical species known from eastern Africa to Southeast Asia and Hawaii, most species of the genus are limited to East and Southeast Asia and the Himalayas in their distribution (He & Snider 2000). According to He & Snider (2000) and other reports, S. perrottetii is the most widely distributed species in the genus, ranging

52 from Sri Lanka, southwestern India, southwestern China, , Thailand, , , Malaysia, Singapore, , Taiwan and Japan. The northern limit of its species distribution is the central part of Honshu, Japan, where snow falls in winter (Kiguchi et al. 2006b); while the southern limit is extended to New Guinea (He & Snider 2000). Higuchi (2014) reported the species from the Hawaii Islands (Hawaii Island), but recently this record was revealed to S. chrysobasilatus, a species endemic to the Hawaii Islands (Kim et al. 2020). Symphyodon perrottetii is characterized by sparsely branched secondary stems, which grow slightly to be erect or long-pendulous from the substrate, weak differentiation between stem and branch leaves, acute leaf apices, serrate upper leaf margins, short and double costae usually reaching less than 1/2 of the leaf length, and weakly prorate laminal cells at the upper cell ends (sometimes smooth). In general, a distinct transverse undulation of the upper lamina is a good feature to identify it in the field. However, this species shows a great diversity in morphological features, which may explain why it has been commonly reported in wide distribution ranges. Indeed, it is often difficult to know the species identity solely based on their morphology. Among the species of Symphyodon, it has known that there is remarkable variation in plant sizes, branching patterns, leaf shapes and lengths, extent of serration at upper leaf margins, and costal lengths. The habit variation is also obvious, with some pendulous from tree trunks and others shortly spread horizontally. The degree of proration of laminal cells is also quite variable too, ranging from indistinct to conspicuous, sometimes accompanying some spines. The spines of the capsule walls likewise vary from smooth to strongly echinate with spines reaching 200 µm in length. With the recent development of applying molecular phylogenetic analyses, many studies have shown that some species previously treated as a single species are actually multi-species complex (Heinrichs et al. 2010; Akiyama 2019; Bakalin et al. 2020). In this study, we have re-examined the Asian Symphyodon species, particularly paying attention to S. perrottetii because of its variation and wide distributional range as described above. Morphological variations and molecular features are analyzed using living samples collected from field work in Japan, Taiwan, China, Vietnam, Thailand, Malaysia, Indonesia and the Philippines, as well as using herbarium material on loan. The results show that the commonly recognized S. perrottetii in the region consists multiple species, including several monophyletic clades, and that each of these clades is well defined morphologically. Furthermore, the phylogenetic position of the genus Rheoshevockia, which is considered to be closely related to Symphyodon, was also analyzed.

Materials and Methods Taxon sampling We analyzed samples identified as Symphyodon perrottetii based on morphological characteristics, as well as other species of the genus. The morphological characteristics of S. perrottetii include ovate-lanceolate leaves with a more or less undulate upper lamina, acute apices with serration in the upper lamina, double costae, and weak proration of laminal cells. We analyzed as many samples as possible, especially from Japan to know distributional patterns of cryptic species.

53 All the 15 species of Symphyodon recognized by He & Snider (2000) were examined morphologically (including type specimens) in this research as well as two additional species, S. chrysobasilatus and S. leiocarpus, which were published after their revision. In addition, we examined type specimens of some species, such as S. merrillii Broth. and Glossadelphus recurvomarginatus Dixon & Sakurai which have been considered as synonyms of S. perrottetii. The following 16 taxa were also analyzed by their DNA features; S. asper (＝S. perrottetii 4 in Fig. 1), S. chrysobasilatus, S. copelandii Broth. (＝S. perrottetii 3), S. complanatus Dixon, S. erinaceus (Mitt.) A.Jaeger, S. erraticus (Mitt.) A.Jaeger, S. gollanioides Nog. (＝S. perrottetii 2), S. imbricatifolius (Mitt.) S.P.Churchill, S. leiocarpus, S. longispinosus, sp. nov. (＝S. perrottetii 7), S. pygmaeus, S. rheophilus, sp. nov. (＝S. perrottetii 5), S. scaber (Tixier) S.He & Snider, and S. scabrisetus Hampe, as well as S. merrillii (＝S. perrottetii 6) and G. recurvomarginatus (＝S. perrottetii 8). These 16 species include 10 of the 15 species recognized by He & Snider (2000). While, no samples of S. angustus (Müll.Hal.) A.Jaeger, S. echinatus (Mitt.) A.Jaeger, S. oblongifolius (Renauld & Cardot) Broth, S. perrottetii s. str., S. weymouthioides Cardot & Thér., and S. yuennanensis Broth. were included in the DNA analysis. As a result, 68 samples of Symphyodon were analyzed. In addition, 6 samples from 5 species of Chaetomitrium spp. (including Chaetomitriopsis), single sample from Dimorphocladon, 2 samples from 2 species of Phyllodon, and 2 samples of Rheoshevockia from the family Symphyodontaceae were also included. For the outgroup, 3 species of Homaliodendron flabellatum (Sm.) M.Fleisch., Taxiphyllum taxirameum (Mitt.) M.Fleisch., and Filibryum ogatae (Broth. & Yasuda) W.Kim & T.Yamag. were used based on previous works (Cox et al. 2010; Kim & Yamaguchi 2016; Ma et al. 2018; Kim et al. 2020).

Examination of herbarium specimens Herbarium specimens borrowed from BM, BO, BORH, CAS, FH, H, HIRO, HSNU, HYO, KYO, L, MAK, MO, NICH, NY, OKAM, OSA, PC, PE, SNP, TNS and UC were used for morphological study in the present survey. As for the specimens deposited in BM and PC, photographs of specimens presented as electrical materials were also used, especially for the type specimens.

DNA extraction, amplification, and sequencing DNA extraction, amplification, sequencing are almost the same as presented in Akiyama et al. (2015) and Akiyama (2017b, 2019) and Akiyama & Odrzykoski (2020). Sequenced regions are rps4, trnL-F, rbcL and ITS1&2. Primers and PCR conditions for rps4, trnL-F, and rbcL are same in Akiyama (2017b). Those for ITS1&2 are ITS5OW (Spagnuolo et al. 1999) and ITS4bryo (Stech 1999) with PCR condition as 94°C (5 min)/ 40°C×[94 (1 min)/ 52°C (1 min)/ 68°C (7 min)]/ 68°C (7 min). Newly obtained nucleotide sequence data reported were deposited in the DDBJ/EMBL/GenBank databases under the accession numbers listed in Appendix. We used MUSCLE (Edgar 2004) to manually align the final consensus sequences, together with previously published sequences from NCBI via internet web browser implemented in MEGA5.2.2 (Tamura et al. 2011). The final data matrix used in the analyses is registered as S28343 in TreeBASE.

54 Phylogenetic analyses Preliminary analyses using rps4, trnL-F, rbcL, ITS1&2 datasets separately suggest the same topology with branches of high supporting values, and thus combined data sets of all four sequences were used for further analyses. Maximum Parsimony (MP) analysis using MEGA5.2.2. (Tamura et al. 2011), and Baysean (BI) analysis using MrBayes ver. 3.2.2 (Ronquist & Huelsenbeck 2003) were used to construct trees. For MP analyses, default settings with TBR branch-swapping with 1000 random addition replicates were used. Non-parametric bootstrapping analyses for MP analysis was employed. This was performed with 500 pseudo-replicates with simple taxon addition in MP analysis. GTR＋G＋I model was used for BI analysis with 1,000,000 generations since it is determined as the optimal model according to the Akaike’s Information Criterion (AIC: Akaike 1973) implemented in MEGA5.2.2; trees were sampled every 1,000 generations. Convergence ESS was assessed by Tracer v.1.7.2 (Rambaut et al. 2018). The first 12,500 trees were removed for the burn-in phrase. Bayesian posterior probability was calculated as node support values.

Results and Discussion The newly obtained sequences comprise 27 species in 5 genera, apart from 3 species in 3 genera of the outgroup (Appendix). A total three most parsimonious trees for MP analysis were obtained with 1090 steps in tree length (CI＝0.556818, RI＝0.832618). Tree topologies from BI and MP analyses did not differ with high support values and thus we used the tree inferred from the BI analyses to represent the results (Fig. 1). In the following part, only the clades supported by both analyses with high supporting values are mentioned. Posterior probabilities of BI analysis (PPBI) and bootstrap values of MP (BSMP) analysis are shown on each branch. Results suggested from the present phylogenetic tree are described and discussed below one by one.

Monophyletic nature of the Symphyodon and systematic position of Rheoshevockia fontana All the samples of Symphyodon spp. and two samples of Rheoshevockia fontana form a well-supported clade (PPBI/BSMP＝1.00/100) and it becomes sister to the clade including all the members of Chaetomitrium, Chaetomitriopsis, Dimorphocladon and Phyllodon (1.00/100). The two samples of Rheoshevockia fontana, which were recently described by Ma et al. (2018) as a distinct genus, were revealed to be the sister group to the clade including all the species of Symphyodon (1.00/100), which supports the opinion to treat it as a separate genus from Symphyodon. This species has an ecological feature as an aquatic rheophyte (Ma et al. 2018), as well as morphological differences from Symphyodon spp. such as fasciculate branching and plane and tight laminal areolation. There is no problem in placing Rheoshevockia as a separate genus within the Symphyodontaceae even if no sporophytes have been found.

Recognition of Symphyodon ovalifolius as sister to all other taxa of Symphyodon Among the Symphyodon perrottetii-like plants shown as S. perrottetii 1–8 in Fig. 1, a total of six samples of S. perrottetii 1 from West Java (Indonesia) and North

55 Figure 1. Molecular phylogenic tree of the Symphyodontaceae based on rbcL, rps4, trnL-F and ITS1&2 sequence data. Supporting values are shown in order of (BIPP/MPBS) beside each branch. Plants previously recognized as S. perrottetii s. l. are shown as S. perrottetii with numbers, and their corresponding species names are indicated on the right.

56 (Malaysia) form a well-supported clade (1.00/100). This taxon is described as a new species, S. ovalifolius H.Akiyama & M.Suleiman below. It is quite notable that the clade is distantly related to all the other members of Symphyodon spp. Overall morphologies of S. ovalifolius are similar to those of other species, but they have long-hanging secondary stems from tree trunks (Fig. 4A & B) similar to S. perrottetii 7, which has been found in central Japan, Taiwan and Vietnam. But it differs from S. perrottetii 7 in branch leaves with more or less round apices and distinctly serration at the upper margins, as well as strong proration on laminal cells of both stem and branch leaves. Therefore, as described in the following taxonomical treatment in detail, S. ovalifolius is proposed to be classified in a new subgenus.

Systematic position of Symphyodon leiocarpus with smooth capsules When Akiyama & Tsubota (2009) described a new species, Symphyodon leiocarpus, from North Thailand, they stressed the smooth capsule walls in contrast to the echinate ones prevailing in the genus and thus proposed to establish a new subgenus Macrothamniopsis H.Akiyama & H.Tsubota. Present analyses using most of the species of Symphyodon reveal that S. leiocarpus forms a monophyletic clade with S. perrottetii 4 (＝S. asper) with echinate capsule walls with a high supporting value (1.00/88). Since closely related genera in the Symphyodontaceae, such as Dimorphocladon borneense Dixon, Phyllodon spp. and most species of Chaetomitrium, have smooth capsule walls, this feature should have been derived by a secondary loss in the genus. Symphyodon angustus has rather lower mammillae on the capsule walls compared to other species of the genus (He & Snider 2000). During examination of herbarium specimens loaned from H, it became evident that S. angustus also had smooth capsule walls [India, West Ghats, South Canara, Kudri-Mukha (Kudremuch), Pfeidere 6444 & 6872; Fig. 2A, smooth capsule in Pfeidere 6444 (H)], although highly mammillose or low echinate ones were often intermingled in the same population [West Ghats, South Canara, Kudri-Mukha (Kudremuch), Pfeidere 6486, 6869 & 6873; Fig. 2B, both highly mammillose (left) and smooth (right) capsules in Pfeiderer 6486 (H)]. Although S. angustus was not included in the present molecular analysis, it is clear that the species is not closely related to S. leiocarpus judging from morphological differences between the two species, such as degree of proration on laminal cells. Therefore, although Akiyama & Tsubota (2009) recognized a subgenus Macrothamniopsis based on the smooth capsule walls of S. leiocarpus, such infrageneric category is unnecessary.

Independency of Hawaiian endemic, Symphyodon chrysobasilaris Glossadelphus chrysobasilaris Broth. was pointed out to be related to the genus Symphyodon by Kim & Yamaguchi (2016), and later Kim et al. (2020) proposed a new combination of Symphyodon chrysobasilaris based on molecular analysis of the sample (U.S.A., Hawaii Islands, Kauai Co., Kawaihau Dist., Shevock, Flynn & Williams 46302, CAS) along with plants collected in the Hawaii Islands and formerly attributed to Glossadelphus. In the present larger-scale treatment, S. chrysobasilaris was confirmed to be a member of the genus as an independent species (Fig. 1).

57 Figure 2. Variation of capsule walls of Symphyodon angustus (Müll.Hal.) A.Jaeger. A: Smooth capsule (Pfeiderer 6444, H). B: Echinate (left) and smooth (right) capsules in a single population (Pfeiderer 6486, H). Scale bars＝0.5 mm for A & B.

Symphyodon pygmaeus widely distributed in Hawaii, Thailand, and East Africa For Symphyodon pygmaeus, two samples from Hawaii and Thailand (Appendix) were analyzed. The present results show the two forming a well-supported clade (Fig. 1, 1.00/100). This result also supports the synonymy of S. sutepensis Tixier from northern Thailand and Glossadelphus abortivapicus Hoe reported from Hawaii (He & Snider 2000). The type specimen of S. pygmaeus from East Africa (Homalia pygmaea Broth., Madagascar, 1890, Chenagor s.n.; isolectotype in H!) is smaller in plant body size and has smaller leaves than specimens from Thailand and Hawaii. In the course of examination of a number of herbarium specimens, however, plant size, leaf size and leaf shape are often quite variable in Symphyodon and thus cannot be reliable features to separate species in the genus. Although S. pygmaeus has a pantropical and quite wide geographical distribution, it seems appropriate to treat it as single species as already pointed out by He & Snider (2000).

58 Symphyodon imbricatifolius, the sole member of the genus in the New World Most species of the genus Symphyodon are distributed mainly in Asia, but only S. imbricatifolius (Mitt.) S.P.Churchill is distributed in South and Central America (He & Snider 2000). In the present study, three samples of S. imbricatifolius are examined and they form a well-supported monophyletic clade (1.00/98) and it is again resolved as a member of the genus. A similar distribution pattern to that found in the genus Symphyodon has previously been reported for the genera Aptychella (Broth.) Herzog (Pylaisiadelphaceae) and Neolindbergia M.Fleisch. (Pterobryaceae); most of their members are distributed in East and Southeast Asia, but A. proligera (Broth.) Herzog distributed in Central and South America (Akiyama et al. 2015) and N. flagellifera Thér. distributed in Cuba (Akiyama et al. 1991). These cases might be good examples of long-distance distribution across different continents in .

Hidden diversity formerly recognized as Symphyodon perrottetii s. l. in East Asia, especially in Japan, Taiwan and southwestern China Plants from East and Southeast Asia that can be identified as Symphyodon perrottetii according to He & Snider (2000), He & Lin (2002) and Kiguchi et al. (2006a, 2006b) were analyzed. Molecular phylogenetic analysis showed that there were at least seven separate monophyletic clades, each labelled as S. perrottetii 2–8 (Fig. 1). Each of Symphyodon perrottetii 2, 3, 4 and 5 formed separately located clades, respectively. Comparison with types and other authentic specimens and morphological studies indicate that S. perrottetii 2 (plants growing on boulders; laminal cells with strong proration) is S. gollanioides Nog., S. perrottetii 3 (plants growing on rocks; laminal cells with strong proration) is S. copelandii Broth., S. perrottetii 4 (plants growing on tree trunk; laminal cells with strong proration) is S. asper, and S. perrottetii 5 (plants growing along streambeds; laminal cells with strong proration) is a new species endemic to Japan described as S. rheophilus below. These four species have been found in inland localities in the Japanese archipelago, such as along deep ravines away from coastal areas. In addition, each of their localities is geographically distantly isolated from each other (Fig. 3B). A total of five specimens of Symphyodon copelandii (as S. perrottetii 3 in Fig. 1) form a clade with high support value (1.00/99). In Japan, this species is strongly restricted to limestone areas of Chichibu District, central Japan. The type locality of S. copelandii is Panay Island (central Philippines), and the Japanese localities of the samples analyzed here are so remote from the type locality. Thus, further molecular analysis comparing plants of Philippines and Japan are needed for more accurate comparison. A total of eight samples of S. asper from Japan and Taiwan (as S. perrottetii 4 in Fig. 1) formed a well-supported clade (1.00/100) and it was sister to the clade of two samples of S. leiocarpus. A total of nine samples of S. rheophilus (as S. perrottetii 5 in Fig. 1) widely distributed in western Japan (Fig. 3B) forms a well-supported clade (0.94/100). In addition, habitat observations of this species revealed that it grows on boulders and/or rocky cliffs in rheophytic zones along streams and small rivers in dense evergreen forests. Rheophytic habitat has been reported in Rheoshevockia, Phyllodon, and Dimorphocladon in the Symphyodontaceae, and

59 Figure 3. Geographical distribution of Symphyodon. A: Four species of S. perrottetii complex (S. perrottetii s. str., S. merrillii, S. longispinosus and S. recurvomarginatus) along with S. yuennanensis. B: Five species (S. asper, S. copelandii, S. gollanioides, S. recurvomarginatus and S. rheophilus) in Japan.

60 thus such a habitat preference can be regarded as secondarily derived in Symphyodon, in which most species are terrestrial. On the other hand, plants designated as Symphyodon perrottetii 6, 7, and 8 (Fig. 1) formed a moderately supported clade (1.00/67). These plants share such morphological features as 1) distinctive undulation on the upper portion of leaves, 2) short double costae, 3) laminal cells with no or weak proration, and 4) serration of upper leaf margins. According to comparison with types and other authentic specimens, S. perrottetii 6 was identified as S. merrillii, which had been regarded as a synonym of S. perrottetii s. str. (He & Snider 2000), S. perrottetii 7 as representing a new species (S. longispinosus below), and S. perrottetii 8 as S. recurvomarginatus (≡Glossadelphus recurvomarginatus), which was also long regarded as a synonym of S. perrottetii s. str. (Ando & Seki 1962; He & Snider 2000). A total of four samples of S. merrillii from Taichung (Taiwan), Mindanao (Philippines), and Flores (Indonesia) forms a moderately supported clade (0.98/62). Morphological examination of type specimens and illustrations suggest the plants should be identified not as S. perrottetii s. str. but S. merrillii, because S. perrottetii s. str. is revealed to have much larger leaves than S. merrillii (Figs. 9 & 10) and these two have different distributional patterns; the former in continental Asia and Sri Lanka and the latter confined to the Malesia region along with a single locality in Laos (Fig. 3A). However, the present molecular phylogenetic analysis did not include DNA samples attributed to S. perrottetii s. str. from the former areas, and thus further study is highly expected to confirm the relationship between S. perrottetii s. str. and S. merrillii proposed here. A total four samples of S. perrottetii 7 from western Yunnan (China) forms a well- supported clade (1.00/100). These Chinese plants have been identified as S. perrottetii (He & Snider 2000; He & Lin 2002), but they have leaves differing in size and shape from S. perrottetii s. str. (Fig. 10), and above all they have capsules with much longer spines (reaching 200 µm in height) than those of S. perrottetii s. str. (at most 90 µm in height). Therefore, present molecular phylogenetic analysis and morphological characteristics suggest that they should be treated as a separate species and are named as S. longispinosus below. There is another species previously described from Yunnan, China. It is Symphyodon yuennanensis (Brotherus 1929). This species has the following morphological features and thus is different from S. longispinosus: more slender leaves, obtuse leaf apices, and more distinctive proration on laminal cells (Fig. 5). However, He & Snider (2000) wrote that leaves of the type specimen of S. yuennanensis were narrowly round at apices, which is not the case as described in the taxonomical treatment below. A total of 14 specimens of Symphyodon perrottetii 8 from western Japan and Tam Dao National Park, north Vietnam form a well-supported clade (1.00/98). Their leaves are similar to S. perrottetii s. str. especially in the distinct undulation of the upper lamina, but are much smaller in size and serration at upper leaf margins is less distinct. In Japan, this species is most widely distributed along the Pacific coast as compared to the other four Japanese species (Fig. 3B) and thus has been repeatedly cited as S. perrottetii in much of the floristic literature (e.g., Ando & Seki 1962; Iwatsuki 1970, 2001; Deguchi 1987; Higuchi & Nishimura 2001). The oldest name applied to this species is Glossadelphus recurvomarginatus Dixon & Sakurai, and thus a new combination under Symphyodon is needed.

61 Unexpected species richness found within single population in Japan By analyzing multiple samples from a single locality, we found that multiple species coexisted in a very small area (Fig. 1, indicated as Symphyodon perrottetii 2, 4, 5, and 8); examples are 1) Todoroki Valley, Tokushima Pref., Japan (three species: S. rheophilus, S. asper and S. recurvomarginatus), 2) Mt. Ichifusa, Kumamoto Pref., Japan, and Inotani Valley, Nichinan, Miyazaki Pref., Japan (two species: S. asper and S. recurvomarginatus), 3) Osuzu Valley, Miyazaki Pref., Japan (two species: S. rheophilus and S. recurvomarginatus, 4) Miyanotani Valley, Mie Pref., Japan (two species: S. gollanioides and S. rheophilus). At the same time, these species show habitat differences; S. rheophilus forms thin colonies on rocks and boulders at or beside streambeds, while S. asper and S. gollanioides form colonies on shrub trunks and boulders in riparian thickets and S. recurvomarginatus on moist windy mountain ridges. This is a good example of the fact that species richness has been overlooked in bryophytes with simple morphology, and suggests that further studies in other bryophyte groups are desirable for an accurate understanding of the diversity in mosses.

Taxonomy The key to the species below treats a total of 22 species of the genus Symphyodon, among which 15 species are included in the phylogenetic tree (Fig. 1). A total of nine species are distributed in Japan and its neighboring countries (S. asper, S. copelandii, S. gollanioides, S. longispinosus, S. merrillii, S. perrottetii s. str., S. recurvomarginatus, S. rheophilus and S. yuennanensis). They are treated in detail in the following section along with newly described S. ovalifolius in Malaysia and Indonesia.

Symphyodon Mont., Ann. Sci. Nat., Bot., sér. 2, 16: 279. 1841.

Key to the species of Symphyodon The following key was modified from He & Snider (2000) and He & Lin (2002); species treated in the present paper are shown in bold with numerical numbers. 1. Distributed in Central and South America ······································ S. imbricatifolius 1. Distributed in Asia, Pacific, and Africa ····························································· 2 2. Distributed in the Hawaiian Islands ································································ 3 2. Distributed in Asia and Africa ······································································· 4 3. Leaves with widely rounded or truncate apices, crenulate at upper margins ··· S. pygmaeus p.p. 3. Leaves acute, almost entire ······················································S. chrysobasilatus 4. Plants long-pendent, reaching 20 cm in length, irregularly and sparsely branched; leaves imbricate, deeply concave above (capsules cylindrical, highly echinate) ····················· 5 4. Plants variable (rarely long-pendent in S. ovalifolius and S. merrillii); leaves appressed or spreading, not imbricate, plane or weakly concave above ······································· 6 5. Leaf apices serrate········································································ S. erinaceus 5. Leaf apices crenulate to entire ·················································· S. weymouthioides 6. Some leaves with rounded to truncate apices in the same stems ······························· 7 6. Leaf apices never rounded to truncate, always obtuse, mucronate or acute ··················· 9

62 7. Leaves widely ligulate, rounded to truncate at apices ························· S. pygmaeus p.p. 7. Some leaves rounded, others widely obtuse or acute ············································· 8 8. Leaves always rounded at apices ··················································· S. complanatus 8. Some leaves round at apices, but others obtuse or acute ············ S. scaber＋S. scabrisetus 9. Leaves widely obovate, obtuse (Costa mostly reaching half of leaf length; exothecial spines long, reaching 150 µm in length; distributed in SW China, north Thailand) ······················································································· 2. S. gollanioides 9. Leaves ovate, lanceolate, or acute-mucronate ····················································10 10. Laminal cells of branch leaves moderately to strongly prorate, often accompanying short spines on back ························································································11 10. Laminal cells of branch leaves smooth or weakly prorate ······································15 11. Branch leaves obovate, strongly dentate at apices ······························ 1. S. ovalifolius 11. Branch leaves ovate, moderately serrate at apices ···············································12 12. Plants slender, filamentous; leaves of secondary stems smaller, mostly less than 1.2 mm long ·····················································································5. S. copelandii 12. Plants teretely or complanately foliated, but not filamentous; leaves of secondary stems longer, mostly longer than 1.2 mm long ··························································13 13. Plants rheophytic; leaf apices widely obtuse; restricted to southwestern Japan ·························································································· 3. S. rheophilus 13. Plants terrestrial; leaf apices acute to acuminate; distributed in India and Nepal ···········14 14. Capsule walls always spinose; distributed in northern India and Nepal ·········································································· S. erraticus＋S. oblongifolius 14. Capsule walls smooth, mammillose or low echinate; distributed in southwestern India ································································································ S. angustus 15. Plants growing in valleys; mostly saxicolous; leaves of secondary stems usually asymmetrical; laminal cells distinctly prorate. (Weakly prorate or smooth in Japanese and Taiwanese plants) ············································ 4. S. asper (including S. echinatus) 15. Plants mostly corticolous; leaves of secondary stems always symmetrical ··················16 16. Laminal cells distinctly prorate. (Stem and branch leaves clearly differentiated in size and shape; known from Yunnan and Guizhou, China) ························ 6-4. S. yuennanensis 16. Laminal cells weakly prorate to smooth ··························································17 17. Plants irregularly and infrequently branched; stem leaves usually longer than 2 mm long, more than 0.8 mm wide ··················································· 6-1. S. perrottetii s. str. 17. Plants more or less pinnately branched; stem leaves less than 2 mm, mostly less than 1.5 mm long, less than 0.6 mm width ·····························································18 18. Plants regularly pinnately branched; leaves, ovate, widely spreading; spines of exothecial cells to 200 µm in length; endemic to Yunnan, China ···················· 6-3. S. longispinosus 18. Plants irregularly pinnately branched; leaves mostly narrowly lanceolate, spreading to patent; spines of exothecial cells less than 75 µm in length. (Tips of secondary stems sometimes becoming flagelliform and long-pendent) ···········································19 19. Laminal cells of branch leaves weakly prorate; costae 1/4–1/3 of leaf length ···6-2. S. merrillii 19. Laminal cells of branch leaves smooth; costae 1/6–1/4 of leaf length ············································································· 6-5. S. recurvomarginatus

63 I. Symphyodon subg. Parasymphyodon H.Akiyama & M.Suleiman, subg. nov.

Diagnosis: Similar to subg. Symphyodon, but differ in branch leaves with deep serration at apices, forming a distinct clade separate from all the other taxa in the genus. Type species: S. ovalifolius H.Akiyama & M.Suleiman. Species included: S. ovalifolius.

1. Symphyodon ovalifolius H.Akiyama & M.Suleiman, sp. nov. Fig. 4

Diagnosis: Similar to Symphyodon merrillii, but secondary stems pendent, sparsely branched, reaching 15 cm in length, branch leaves obovoid, roundly obtuse, with distinct serration at apices and laminal cells prorate at upper ends, often intermingled with low spines. Type: MALAYSIA. Sabah, , Liwagu trail, 1,600 m elev., Akiyama & Suleiman 23466 (holotype BORH, isotypes BM, H, HYO, NY & SNP) Etymology: The species was named after its ovate leaves. Description: Plants slender, yellowish green, resembling Symphyodon erinaceus or S. copelandii in general shape. Primary stems short, attached to substrate. Secondary stems soft, often long-pendent, reaching 15 cm in length, sometimes short (ca. 2 cm in length), horizontally spreading, pinnately and sparsely branched, loosely and teretely foliated. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Secondary stem leaves and branch leaves not so much differentiated in size and shape. Stem leaves ovate-lanceolate, acute, 1.3–1.4 mm long, 0.4–0.45 mm wide; costae double, 2/5 to 1/3 of leaf length; alar differentiated with quadrate to short rectangular cells in small numbers. Branch leaves ovate to elliptic, acute to obtuse, 0.9–1.0 mm long, 0.35–0.4 mm wide; costae double, 1/4 to 1/2 of leaf length; alar differentiated with quadrate to short rectangular cells. Leaves of long-pendent secondary stems and those of shortly spreading secondary stems different in shape and degree of proration on laminal cells; leaves oblong to ovate-oblong, widely acute or acute, with weak undulation at upper lamina, distinct serration at upper margins, weak proration on laminal cells in the former case, while ligulate or oblong-lanceolate, mostly obtuse, distinctive prorate on laminal cells (often proration becomes short spines) in the latter case. Sexual organs and sporophytes unknown. Distinguishing features: 1) Secondary stems often pendulous from tree trunks and shrub branches, 2) stem and branch leaves scarcely differentiated in size and shape, 3) branch leaves obovate, narrow at base, distinctly serrate at apices, and 4) laminal cells with distinct proration often becoming short spines. Habitat: Hanging from tree trunks and shrub branches in filtered light at floor of tropical upper montane developed in a valley. Distribution: Malaysia (Sabah) and Indonesia (Java and Flores). Paratypes: MALAYSIA. Sabah, Kinabalu Park, Liwagu trail, 1,600–1,700 m elev., Akiyama & Suleiman 23433, 23434, 23439, 23443, 23463, 23464 and 23467 (all BORH, HYO, SNP). INDONESIA. West Java, Cibodas, G. Gedeh, 1,540 m elev., Akiyama & Haerida 24856 (BO, HYO); ibid., 1,570 m elev., Akiyama & Haerida 24858 (BO, HYO). Flores, above Ruteng: at Poco Gurung in , ca. 1,700 m elev., Schmutz 5278C (L); ibid., Prov.

64 Figure 4. Symphyodon ovalifolius H.Akiyama & M.Suleiman. A: Plant with hanging secondary stems. B: Close-up of secondary stems. C–F: Stem leaves. G–I: Branch leaves. J & K: Leaf apices. L: Upper laminal cells with high proration. M & N: Leaf bases. O & P: Median laminal cells with distinct proration. A–D, G–H, K, M, N & O from Akiyama & Suleiman 23466 (holotype, BORH), E, F, I, J & P from Akiyama & Haerida 24856 (HYO). Scale bars＝1 cm for A, 2 mm for B, 0.1 mm for C–I & M＝ 20 µm, 10 µm for others. 65 Manggarai, Ruteng, Poco Tador Walok, between lumberjack’s camp and the summit, 2,050– 2,130 m elev., Schmutz 6826 (L). Taxonomical note: He & Snider (2000) reported Symphyodon copelandii from Kinabalu Parks [Borneo Isl., Sabah, Kamborangah, Holttum 25680 (BM)]. Judging from the locality of the specimen, it seems as if this is the new species described here, even though the authors did not examine the specimen. It is true that S. ovalifolius and S. copelandii are similar to each other in having long-pendent secondary stems. The specimen formerly identified as Symphyodon copelandii Broth. collected in Flores Island (Ruteng, Veldkamp 7112, L!; may be transcription error of 7121) by He & Snider (2000) was revealed to be S. merrillii Broth., which is treated as a distinct species here. On the other hand, the two specimens collected in Flores Island cited above have similar morphological features to the typical plants of S. ovalifolius in Sabah and West Java but have deeply concave leaves. Symphyodon perrottetii var. robusta M.Fleisch., nom. nud. was reported from West Java. Plants of the variety have long pendent secondary stems hanging from branches of small shrubs. It looks similar to S. ovalifolius, S. copelandii, or S. recurvomarginatus in general appearance, but a detailed examination of specimens clearly suggests that it is S. merrillii.

II. Symphyodon subg. Symphyodon Mont., Ann. Sci. Nat., Bot., sér. 2, 16: 279. 1841. Type species: Symphyodon perrottetii Mont. ＝Symphyodon subg. Macrothamniopsis H.Akiyama & H.Tsubota, Acta Phytotax. Geobot. 60: 93. 2009; syn. nov. Type species: Symphyodon leiocarpus H.Akiyama & H.Tsubota.

2. Symphyodon gollanioides Nog., Lindbergia 1: 180. 1972. Figs. 3B & 5A–D Type: THAILAND. Payap: Kun Kawng San, on twig, 1,700 m elev., Hansen et al. 11428 (holotype C, isotype NICH!)

Description: Plants yellowish-green, turning yellow in older specimens, not shinny. Primary stems shortly prostrate on substrate. Secondary stems dense and more or less complanately foliate. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Secondary stem leaves to 1.6 mm long, lanceolate to ovate, symmetric or slightly asymmetric, plane and not undulate, widely acute, weakly to moderately serrate at upper margins; laminal cells linear, weakly prorate to distinctly prorate, thin- to weakly thick-walled; costae double, reaching 1/3 to 2/5 of leaf length; alar cells well-developed, rectangular, not thick-walled. Branch leaves 0.9–1.2 mm long, narrowly ovate to obovate, more or less asymmetric, plane, often narrowed at base, acute or obtuse, densely serrate at upper margins; laminal cells weakly prorate to distinctly prorate; costae double, 2/5 to 1/2 of leaf length; alar cells weakly developed, quadrate, not thick-walled. Dioicous. Inner perichaetial leaves abruptly narrowed above; costa double but indistinct. Perigonia not observed. Setae 13–18 mm long, mammillose above, smooth below. Capsules slightly inclined, oblong-ovoid, 1.3–2.2 mm long, with dense and long spines reaching 150 µm in length; annuli differentiated. Opercula conic, ca. 1.2–1.5 mm long. Peristome

66 Figure 5. Symphyodon gollanioides Nog. (A–K) and S. yuennanensis Broth. (L–O). A, B & L: Plants in dry condition. C: Close-up of secondary stems and branches. D–E & M: Secondary stem leaves. F–H: Branch leaves. I: Alar region of secondary stem leaves. J: Median laminal cells. K, N & O: Upper part of lamina. A from Hansen et al. 11428 (isotype, NICH), B–K from Akiyama 21857 (HYO), L–O from Handel-Mazzetti 9473 (holotype, H-BR). Scale bars＝1 cm for A, B & K, 0.5 cm for C, 100 µm for D–H & K, 10 µm H–J, 20 µm for M & N.

67 double. Exostome teeth linear-lanceolate, to 700 µm in height, densely papillose. Endostome segments linear, ca. 2/3 length of exostome teeth; basal membranes low, 60–80 µm in height. Spores spherical, minutely papillose, ca. 20 µm in diameter. Distinguishing features: 1) Ovate to obovate stem and branch leaves with somewhat narrowed bases, 2) often only weakly prorate laminal cells, 3) rather long costae, 4) weak differentiation of alar cells in branch leaves, and 5) highly echinate capsules with long spines, reaching 150 µm in length. Habitat: Forming a thin turf on boulders in a deep ravine in mixed forest. Distribution: Thailand, India, and Japan. Other specimens examined: INDIA. Sikkim, Kurseong, Sepoydura forest, 1,800 m elev., Decoly & Schaul s.n. (H). JAPAN. Mie Pref., Matsusaka-shi, Iitaka-cho, Hachisu, 34°20′35″N, 136°08′46″E, 485 m elev., Akiyama 21857 (HYO). Taxonomical notes: The figures (Figs. 187–194) in He & Snider (2000) based on the holotype (Hansen et al. 11428, C) differs from the figure (Fig. 4) in Noguchi (1972) in the following features; 1) broadly ovate leaves, 2) distinctly thick-walled cells of upper, median and alar cells, and 3) highly echinate capsule walls. Observations based on the isotype specimen borrowed from NICH confirmed that leaves are often slightly asymmetrical, narrowly lanceolate or ovate, acute or broadly acuminate, with more or less evenly and slightly thick-walled laminal cells, and alar cells not porose, as described by Noguchi (1972). In addition, laminal cells of the plants from Japan and India (Sikkim) are not particularly thick-walled. While the spines on the capsule wall were illustrated too low in Noguchi’s figure. They are actually about 150 µm in height, which is more accurate in He & Snider’s (2000, Fig. 191). The specimen from India (Decoly & Schaul s.n.) was cited as both Symphyodon perrottetii and S. gollanioides in He & Snider (2000). Examining the specimens, morphological features fit well with those of S. gollanioides.

3. Symphyodon rheophilus H.Akiyama, sp. nov. Figs. 3B & 6

Diagnosis: Similar to Symphyodon merrillii, but plants more or less complanately foliate, stem and branch leaves acute to obtuse, laminal cells strongly prorate, habitat confined to rheophytic zones along small rivers. Type: JAPAN. Miyazaki Pref., Tsuno-cho, Osuzu-keikoku, in the vicinity of Yagen Waterfall, 500 m elev., Akiyama 22870 (holotype HYO, isotypes BM, H, NICH and NY) Etymology: Named after the habitat preference for rheophytic zones. Description: Plants green, not glossy. Primary stems pinnately branched, prostrate on substrate. Secondary stems prostrate, irregularly branched, to 4 cm long, densely and ± complanately foliate. Paraphyllia and pseudoparaphyllia absent. Branches less than 1 cm long. Asexual propagules absent. Stem and branch leaves of ascending stems differentiated. Stem leaves lanceolate to ovate lanceolate, acute to widely acute, 1.2–1.7 mm long, loosely undulate above, crenulate at upper margins; laminal cells fusiform to linear, prorate; costae double, 1/5–1/4 (–1/3) of leaf length; alar cells differentiated, quadrate to short rectangular. Branch leaves ovate, widely acute to obtuse 0.8–1.1 mm long, minutely serrate to crenulate at upper

68 Figure 6. Symphyodon rheophilus H.Akiyama. A: Plant. B: Secondary stems and branches. C: Close-up of branches. D–F: Secondary stem branches. G & H: Branch leaves. I–J: Apices of secondary stem leaves. K & L: Median lamina cells of secondary stem leaves. M: Median laminal cells of branch leaf. N & O: Alar regions of secondary stem leaves. All from Akiyama 22870 (holotype, HYO). Scale bars＝1 cm for A, 5 mm for B, 2 mm for C, 100 µm for D–H, 10 µm for I & J, 10 µm for K–O.

69 margins, laminal cells prorate, often with short spines; costae 1/4–1/3 of leaf length; alar cells differentiated, quadrate. Sexual organs and sporophytes unknown. Distinguishing features: 1) Habitat preference for rheophytic zones in warm temperate, deciduous forests, 2) complanately foliate secondary stems and branches, 3) weak serration at upper leaf margins, 4) acute to obtuse leaf apices, and 5) strong proration on laminal cells. Habitat: According to the author’s observations, this species grows on boulders in upper rheophytic zones along small rivers running through dense lower montane forests. Distribution: Endemic to southwestern Japan. Paratypes: JAPAN. Mie Pref., Matsusaka-shi, Iitaka-cho, Hachisu, Miyanotani valley, 530 m elev., Akiyama 21855, 21858 and 21861 (HYO); ibid., 500 m elev., Akiyama 24986, 25000 & 25001 (HYO); ibid., 570 m elev., Kiguchi & Tateishi 40734 (HYO ex herb. Kiguchi). Tokushima Pref., Kaiyo-cho, Todoroki-Kujyuku-taki Falls, Nabewari-taki fall, 450 m elev., Nishimura 13225 (HYO, OKAM); ibid., Nabewari-taki fall, 495 m elev., Tateishi 33138 (HIRO, HYO). Miyazaki Pref., Tsuno-cho, Osuzu-keikoku valley, Akiyama 22875, 22892 & 26028 (HYO). Taxonomical note: This species is restricted to boulders in upper rheophytic zones along small rivers and is a good examples of rheophytic species in mosses. In the Japanese archipelago, which is the northern limit of distribution of Symphyodon, it is likely that an ancestral terrestrial species expanded its habitat into rheophytic environments as in the case of another rheophyte moss in the Symphyodontaceae, Rheoshevockia fontana.

4. Symphyodon asper (Mitt.) A.Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1876–77: 296. 1878. Figs. 3B & 7 ≡Stereodon asper Mitt., J. Proc. Linn. Soc. Bot. Suppl. 1: 110. 1859. Type: INDIA. Khasia, Hooker & Thomson 866 (lectotype NY, isolectotypes BM, H!)

Description: Plants shortly prostrate on substrate. Secondary stems 1–3 cm long, bipinnately to tripinnately branched. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Stem and branch leaves not so much differentiated in size and shape. Stem leaves 1.0–1.4 mm long, lanceolate to ovate, often asymmetric and narrowly decurrent at bases, acute, denticulate above; upper laminal cells short-rhomboidal, mostly smooth; median laminal cells linear, indistinctly prorate or almost smooth (especially in Japanese plants); alar cells differentiated, quadrate; costae double, short, 1/4–1/5 of leaf length. Branch leaves 0.6–0.9 mm long, oblong to ovate, often asymmetric and narrowly decurrent at bases, acute or narrowly obtuse, denticulate in upper 2/3 of leaf margins; upper laminal cells short- rhomboidal, weakly prorate; median laminal cells linear, distinctly prorate; alar cells differentiated, quadrate; costae double, short, 1/5–1/6 (rarely less than 1/7 or indistinct) of leaf length. Dioicous. For sexual organs and sporophytes, see He & Snider (2000) since they were not observed among the specimens we examined. Distinguishing features: 1) Bipinnately to tripinnately branched plants, 2) slightly dimorphic leaves in shape and size between stems and branches, 3) often asymmetric leaves with more or less decurrent bases, 4) weak (stem leaves) to distinct (branch leaves) proration on laminal cells, and 5) strongly papillose exostome teeth (He & Snider 2000).

70 Figure 7. Symphyodon asper (Mitt.) A.Jaeger. A: Plants. B: Close-up of secondary stems and branches. C–G: Secondary stem leaves. H & I: Branch leaves. J–L: Leaf apices. M–N: Median laminal cells of secondary stem leaves. O: Median laminal cells of branch leaf. P–Q: Alar regions of secondary stem leaves. C, D, L & N from Akiyama 23254 (HYO), others from Akiyama 22959 (HYO). Scale bars＝ 5 mm for A & B, 100 µm for C–I, 10 µm for J–Q.

71 Habitat: Growing on boulders along deep ravines in evergreen forests or shortly hanging from small shrub trunks and branches at more or less mossy montane forests. Distribution: Japan, Taiwan, Thailand; reports from Vietnam, and India, see He & Snider (2000). Representative specimens examined: JAPAN. Kochi Pref., Aki-gun, Umaji-mura, Ganmaki Valley, Deguchi 27319 (KOCH, NICH); Susaki-shi, Mt. Honokawa, Kuwanomata Valley, ca. 700 m elev., Iwatsuki s.n. (NICH); Kitagawa-mura, Takeyashiki, Ogawa River, 435 m elev., Tateishi 35324 (HIRO). Tokushima Pref., Kaiyo-cho, Todoroki-Kujyuku-taki Falls, trail to a shrine, 255 m elev., Tateishi 33116 (HIRO); ibid., 240 m elev., Nishimura 13212 (OKAM). Kumamoto Pref., Mizukami-mura, Mt. Ichifusa, en route from the entrance to Ichifusa Shrine, 790 m elev., Akiyama 23594 (HYO). Miyazaki Pref., Nichinan-shi, Inotani, 300 m elev., 31°40′N, 131°11′E, Akiyama 22959 (HYO). Kagoshima Pref., Yakushima Isl., near Yodogo mountain hut, ca. 1420 m elev., Suzuki 11522 [NICH; quoted as S. echinatus in Suzuki (2017)]; ibid., between Onoaida to Jyataki Fall, 150–500 m elev., Iwatsuki et al. M1177 (NICH). TAIWAN. Miaoli Co., Tai’an Township, Hshuepa National Park, Syuejin Recreation Area, along 1st observing trails, 1,850 m elev., 24°25′N, 121°00′E, Akiyama 23149, 23231 and 23254 (HYO). THAILAND. Chiang Mai, Doi Inthanon, ca. 2,250 m elev., Akiyama et al. 1250 & 1283 (both HYO). Taxonomical notes: Plants growing in Japan and Taiwan differ from the typical plants: they have bipinnately or tripinnately branching pattern but stem and branch leaves are mostly similar in size and shape. In addition, they often have weak proration on laminal cells, especially for those of stem leaves, and short costae compared to plants occurring in South Asia. He & Snider (2000) pointed out that leaf apices ending in three cells is one of the good distinguishing features of Symphyodon asper. However, in terms of the examined plants of East and Southeast Asia including Japan and Taiwan, the numbers of cells forming the leaf apices are not stable within species or variable even within the plants from the same specimen packet. In addition, He & Snider (2000) wrote that Symphyodon echinatus was able to be distinguished from S. asper by the symmetric leaves with strongly decurrent bases, leaf apices ending in one cell, and finely papillose exostome teeth of the latter species. According to my observations, Symphyodon asper and S. echinatus share a number of morphological features and they are scarcely distinguished from one another except for asymmetric leaves and weaker proration on laminal cells in S. asper. Suzuki (2017) reported Symphyodon echinatus from Japan (Yakushima Island, Suzuki 11522, NICH) based on a specimen that includes only a small fragment of a single stem. Judging from the rather poor proration of laminal cells and weakly asymmetric leaf shape, it seems appropriate to identify the specimen not as S. echinatus, but as S. asper as treated in the “Specimens examined” above. However, their inter-relationship needs further study in the future as pointed above.

5. Symphyodon copelandii Broth., Philippine J. Sci. 31: 291. 1926. Figs. 3B & 8 Type: PHILIPPINES. Panay, Benguet Prov., Pauai, Copeland 1337 (holotype H-BR!, isotype FH)

72 Figure 8. Symphyodon copelandii Broth. A & C: Plants. B & D: Close-up of secondary stems. E–F: Stem leaves. G–J: Branch leaves. K & L: Leaf apices of branch leaves. M: Basal part of branch leaf. N: Lower median laminal cells. O: Upper laminal cells. A & B from Copeland 1337 (holotype, H-BR), others from Akiyama 23014 (HYO). Scale bars＝1 cm for A & C, 2.5 mm for B & D, 100 µm for E–J, 10 µm for I–O.

73 Description: Plants dull green, filamentous and thread-like, irregularly or more or less pinnately branched, creeping on substrate. Stems thin, ca. 0.15 mm wide (0.5–0.8 mm wide including leaves), teretely or slightly complanately foliate; leaves appressed to stems or narrowly spreading. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Stem and branch leaves more or less differentiated. Stem leaves oblong-ovate, acute, 0.5–1.2 mm long, 0.2–0.45 mm wide, flat, narrowly decurrent at both sides at base; margins densely serrulate above, almost entire at base; costae double, 1/3–1/5 of leaf length; upper laminal cells fusiform, distinctly prorate at upper ends; median laminal cells distinctly prorate, basal laminal cells narrow; alar cells few, quadrate. Branch leaves narrowly lanceolate, less than 0.5 mm long, ovate; laminal cells linear, strongly prorate, often becoming short spines; costae double, reaching 1/5–1/4 of leaf length. Dioicous. Perichaetial leaves more or less abruptly acuminate and entire or minutely serrate at apices, ecostate. Setae ca. 1.5–2.0 cm long, mammillose above, smooth below. Capsule erect, oblong-ovoid, slightly asymmetric, 1.6–1.8 mm long, densely echinate and the spines reaching 95 µm in length. Peristome double. Exostome teeth smooth. Endostome segments linear, smooth. Spores ca. 20 µm in diameter, distinctly papillose. [Sexual condition and sporophyte description are according to Brotherus (1926) and He & Snider (2000).] Distinguishing features: 1) Slender and often filamentous plants, 2) stem and branch leaves not differentiated in shape, but different in size, 3) branch leaves less than 0.5 mm in length, 3) strongly prorate laminal cells in stem and branch leaves, and 4) short costae, 1/5–1/4 of leaf length. Habitat: According to our observations on Japanese plants, they grow on chart cliffs and boulders in limestone areas, rarely on shrub branches just beside a cliff in filtered light, in a rather moist places along streamlets in a deciduous forests. Distribution: Indonesia (Flores), Philippines (Luzon, Panay), Japan (central Honshu). In Japan, this species is known from the limestone areas in the Chichibu district of the Kanto region under the name of Symphyodon perrottetii (Kiguchi et al. 2006b), where it snows in winter. The distribution area of the species is the highest latitude for the genus Symphyodon, which is mainly distributed in tropical and subtropical countries. Representative specimens examined: JAPAN. Saitama Pref., Chichibu-shi, Nakatsugawa, Gakushu-no-mori, 39°59′13″N, 138°47′14″E, 820 m elev., Akiyama 23014 & 23016 (HYO); Chichibu-shi, Ochigawa river, Nishitani, 1,000 m elev., Nagano 6725 (HYO ex herb. Kiguchi); Ogano-cho, Kawarasawa, above the entrance of Haccho-tunnel, 36°02′24.8″N, 138°49′51.7″E, 1,240 m elev., Akiyama 22992, 22993 & 22994 (HYO); Ogano-cho, east slope of Mt. Futago, 36°04′16.8″N, 138°52′15.4″E, 800 m elev., Kiguchi & Hiraoka 74 (HYO ex herb. Kiguchi). Tokyo Metro., Okutama-cho, Ogawadani, 35°53′13.1″N, 129°01′9.8″E, 1,150 m elev., Kiguchi & Suzuki 50535 (HYO ex herb. Kiguchi). PHILIPPINES. Luzon, Mt. Tabayoc, Lake Ingcolos, 16°42′N, 120°53′E, 2,420–2,450 m elev., Jacobs B205 (H, L); ibid., Reynoso 2182 (MO). INDONESIA. Flores Island, Ruteng, Veldkamp 7112 (L, MO). Taxonomical notes: Sporophytes and sexual organs have not been found from Japanese plants. In addition, the author was not able to find sporophytes in the packet of the holotype deposited in H. He & Snider (2000) reported Symphyodon copelandii from Kamborangah (Borneo), near

74 Liwagu trail in Kinabalu Park, which might be S. ovalifolius where a number of specimens of the species were collected. In addition, they cited another specimen of S. copelandii from Indonesia (Flores, Ruteng, Poco Ranaka, 2,000 m elev., Veldkamp 7121B (L), quoted as 7112), which is treated as S. merrillii in this paper. Symphyodon copelandii has the most slender plants in the genus, and it sometimes becomes filamentous in shaded, darker conditions. In addition, the shape, size and degree of proration of the laminal cells tend to differ between leaves of laterally spreading short stems and long-pendent stems. Only samples from Japan were examined for DNA features of S. copelandii, and no sample from the type locality (Philippines) was included. In other words, the identification of Japanese plants as S. copelandii was based only on morphological similarity. Therefore, detailed relationships between plants of Philippines and Japan will be the subject of future studies using molecular markers.

6. Symphyodon perrottetii and related species Based on the present phylogenetic analysis, the Symphyodon perrottetii and related species are revealed to include S. perrottetii s. str., S. merrillii, S. longispinosus, and S. recurvomarginatus. Although S. merrillii was synonymized under S. perrottetii by He & Snider (2000), its resurrection is proposed here, even though no sample of S. perrottetii s. str. itself was included in the molecular analyses. Their relationship is inferred mostly by their differences in morphological features (Figs. 9 & 10). Symphyodon perrottetii s. str. was originally described on the basis of a specimen collected at Nilghiri Hill, Tamir Nadu, southwestern India. It has larger leaves that are strongly undulate above, deeply serrate at the upper margins and with smooth laminal cells. Capsules are oblong-ovate with spines less than 90 µm in length (Fig. 9E). On the other hand, S. merrillii, whose type locality is the Philippines, is characterized by slightly smaller leaves than those of S. perrottetii and weakly to moderately prorate laminal cells. Capsule shape is almost similar to those of S. perrottetii. Therefore, the relationship between S. perrottetii s. str. and S. merrillii requires further analysis, especially including samples from southwestern India and Sri Lanka.

6-1. Symphyodon perrottetii Mont. s. str., Ann. Sci. Nat., Bot., sér. 2, 16: 279. 1841. Figs. 3A, 9D, E, H, J, K, N, O & 10A–D Type: INDIA. Tamir Nadu, Nilghiri, Perrottet s.n. [holotype PC, isotypes BM, H (?)]

Description: Plants large to robust. Primary stems long, reaching 15 cm in length, densely pinnately branched, prostrate. Secondary stems ascending and irregularly and sparsely branched, to 5 cm long, densely and more or less roundly or slightly complanately foliate. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Branches 0.5–1.0 cm long. Stem and branch leaves not clearly differentiated, patent or slightly spreading when dry and wet. Stem leaves 1.6–2.7 mm long, ovate-lanceolate, scarcely or slightly decurrent at base, acute, serrate at upper margins, often deeply undulate above; upper laminal cells rhomboidal to linear, smooth; median laminal cells linear, scarcely prorate or smooth; alar cells

75 Figure 9. Symphyodon merrillii Broth. (A–C, F–G, I, L–M) and S. perrottetii Mont. s. str. (D–E, H, J, K, N–O). A–B, D: Plants with sporophytes. C & E: Capsules. F, G, J, K: Secondary stem leaves. H & I: Apices of stem leaves. L & N: Alar regions of stem leaves. M & O: Median laminal cells of stem leaves. A–C, F, G, I, L & M from Merrill 6193 (isotype of S. merrillii, H), D, E, H, J, K, N & O from Perrottet 1368 (isotype? of S. perrottetii, H). Scale bars＝5 mm for A, B & D, 1 mm for C & E, 100 µm for F, G, J & K, 10 µm for H, I, & L–O.

76 differentiated, quadrate; costae double, reaching 2/5–1/3 of leaf length. Branch leaves smaller, 1.0–1.2 mm long, ovate-lanceolate to lanceolate, shallowly undulate above, otherwise similar to stem leaves; laminal cells smooth to scarcely prorate; alar cells differentiated, quadrate; costae 2/5–1/3 of leaf length. Calyptrae cucullate, smooth. Dioicous. Perichaetial leaves gradually narrowed to long acumen; laminal cells thick- walled, costae 2–3, reaching 1/4 of leaf length. Setae 12–18 mm long, reddish brown, mammillose above, smooth below. Capsules erect, cylindrical, symmetric, 2.8–3.3 mm long, densely and moderately echinate; spines mostly less than 90 µm in length. Opercula long rostrate, ca. 1.4 mm long. Peristome double. Exostome teeth narrowly lanceolate, ca. 500 µm in height, smooth to finely papillose. Endostome segments reduced, adnate to the exostome teeth at base; basal membranes obscure. Spores spherical, ca. 17 µm in diameter, papillose. [Peristome features after He & Snider (2000)] Distinguishing features: 1) Robust plants that are irregularly and sparsely branched, 2) robust leaves, reaching 2.7 mm in length, 3) deep undulation at upper part of lamina, 4) deep serration at upper leaf margins, 5) almost smooth laminal cells, and 6) cylindrical capsules with spines of moderate length. Habitat: No information available. Distribution: Southwestern India, Sri Lanka, Singapore, and Thailand. Representative specimens examined: INDIA. India orient., Neilghery, Perrottet 1632 (probably isotype, H, PC). Tamil Nadu, Nilghiri Hills, ca. 6 km from Ootacamund on road to Kotagiri, Townsend 73/593 (MO); ibid., Avalanhe Valley between Avanche and Emerald, Townsend 73/682 (MO). SRI LANKA. Pedrutalagala, 2,400 m elev., Binstead 228 (H). Central Prov., Nuwara Eliya, Binstead 316 (H). SINGAPORE. without exact locality, Douin 28 (H). THAILAND. Nakornsrithamarat, Khao Luang, 8°30′N, 99°45′E, 1,250–1,450 m elev., Touw 11671 (L, MO). Taxonomical notes: There is a good illustration of Symphyodon perrottetii in the original description (Montague 1841). It suggests clearly that S. perrottetii has long, wide leaves and low spines on the capsule. High resolution images of holotype and isotypes are deposited in BM and PC and they are quite helpful to recognize such features. As discussed above, plants previously identified as Symphyodon perrottetii from Japan, Taiwan, Philippines and China, are treated as three separate species in this paper; they are S. merrillii, S. longispinosus, and S. recurvomarginatus.

6-2. Symphyodon merrillii Broth., Philippine J. Sci. 2: 341. 1907. Figs. 3A, 9A–C, F, G, I, L, M & 10E–L Type: PHILIPPINES. Mindoro, Mt. Halcon, 1,800 m elev., Merrill 6193 (holotype H!, isotype H!) ＝Symphyodon perrottetii auct. non Mont. sensu Fleisher (1923: 1111, Fig. 186) ＝Symphyodon perrottetii f. robusta M.Fleisch., Musci Fl. Buitenzorg 4: 1668. 1923; nom. nud.

Description: Primary stems prostrate on substrates, densely and pinnately branched. Secondary stems 2–4 cm long, sparsely to densely pinnately branched, rarely becoming

77 Figure 10. Comparison of leaves between Symphyodon perrottetii s. str. (A–D) and S. merrillii (E–M). A: Thailand, Khao Luang, Touw 11671 (MO). B: India, Neilgiri Hills, Perrottet 1632 (H, PC; isotype? of S. perrottetii). C: Sri Lanka, Nuwara Eliya, Binstead 316 (H). D: Singapore, Douin 28 (H). E: Indonesia, Java, Tjibodas, Nyman 361 (H, L). F: Indonesia, Halmahera, Vriese s.n. (H). G: Philippines, Mindanao, Shevock 44824 (CAS). H: Laos, Ban Pha Lam Phen, Tuyama s.n. (NICH). I: Taiwan, Taoyuan Co., Shevock 30525 (CAS). J: Indonesia, Halmahera, Sande Lacoste s.n. (L). K: Indonesia, Seram, Vriese s.n. (H). L: Philippines, Mindoro, Merrill 6193 (isotype of S. merrillii, H). M: Indonesia, Flores, Touw & Snoek 23213 (L). Scale bars＝0.2 mm.

78 slender and long-pendulous at tips, reaching 10 cm in length. Branches short, usually less than 10 mm long. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Stem and branch leaves not much differentiated in size and shape except for those of pendulous parts. Stem leaves narrowly ovate-lanceolate to lanceolate, not decurrent at base, 1.4–2.1 mm long, acute, serrate to serrulate at upper margins, somewhat weakly incurved below, undulate above; laminal cells weakly prorate or sometimes smooth; costae 1/4–1/3 of leaf length; alar cells differentiated quadrate to short rectangular. Branch leaves ovate lanceolate to narrowly lanceolate, not decurrent, 1.2–1.4 mm long, acute, serrulate at upper margins, weakly incurved below; laminal cells rhomboidal to linear, weakly prorate; costa 1/5–1/4 of leaf length; alar cells few, quadrate. Calyptrae cucullate, naked. Dioicous. Perichaetial leaves sheathing at base, serrulate at upper margins. Setae straight, 22–30 mm long, reddish brown, highly mammillose above, smooth below. Capsules cylindrical, upright, 2.5 mm long, 0.6 mm wide, densely echinate, spines less than 75 µm in height. Opercula long, narrowly rostrate, ca. 2 mm in length. Peristome double, pale white. Exostome teeth narrowly lanceolate, minutely papillose. Endostome segments short, linear, keeled; basal membranes low. Spores spherical, ca. 15 µm in diameter, minutely papillose. Distinguishing features: 1) Weakly differentiated leaves between stems and branches, 2) narrowly ovate to linear-lanceolate stem leaves, 1.4–2.1 mm in length, 3) more or less undulate upper lamina, 4) costae 1/4–1/3 of leaf length, 5) weakly prorate laminal cells, and 6) low echination of capsule walls. Habitat: On twigs and trunks of small diameter hardwoods in undergrowth of tropical montane forests, mostly along narrow valleys. Distribution: Taiwan, Laos, Philippines (Luzon, Mindanao, Mindoro), Indonesia (Java, Lombok, Flores, Halmahera, Seram). Papua New Guinea (Central District). Representative specimens examined: TAIWAN. Taoyuan Co., Lalashan National Forest Reserve, 24°42′59″N, 121°26′27″E, 1,600 m elev., Shevock & Yang 30525 (CAS). LAOS. Ban Phu Phao (Ban Pha Lam Phen?), ca. 25 km NW of Phongasavanh, 1,200 m elev., Tuyama s.n. (NICH). PHILIPPINES. Mindanao, Mt. Kitanglad Range Natural Park, midslope of Mt. Dulang-Dulang, 08°05′42.0″N, 124°55′26.6″E, 2,130 m elev., Shevock 44714 (CAS); ibid., 03°06′36.0″N, 124°55′30.0″E, 2,675 m elev., Shevock 44824 (CAS); Bukidnon Prov., Kalatungan, 2,000 m elev., 28 Dec. 2012, Segumpan et al. s.n. (CAS, UC). Luzon, Mt. Pulog, 16°36′N, 120°54′E, 2,600–2,700 m elev., Jacobs B205 (H, L); Mt. Tabayoc, Lake Ingcolos, 16°42′N, 120°53′E, 2,420–2,450 m elev., Jacobs B415 (H, L). INDONESIA. West Java, Pangarango, Nyman 263 (H, L); Gedeh am Gegerbintang an Aesten 1,600–1,700 m, Fleischer s.n. (Exicatae, M. Fleischer, Musc. Archip. Ind. et Polynes. Ser. XI). Lombok, Gunung Rinjani, ascent from Senaro (South of Bayan), ca. 1,950–2,000 m elev., Touw & Snoek 22341 (L). Flores, Manggarai Prov., N. slope of Ngando Napu, South of Ruteng, 1,780 m elev., Touw & Snoek 23213 (L). Halmahera, without exact locality, 1850–1860, Vriese s.n. (H, L). Seram, without exact locality, 1858–1860, Vriese s.n. (H). PAPUA NEW GUINEA. Central Dist., Woitape, on track to Mt. Albert Edward Wharton Ranges, above 2,100 m elev., Robbins 4224 (L). Taxonomical notes: He & Snider (2000) treated Symphyodon merrillii as a synonym of S. perrottetii s. l. It is true that they share weakly differentiated stem and branch leaves,

79 marked undulation at upper lamina, weak proration on upper ends of laminal cells, and low echination on short cylindrical capsules. The latter, however, can be distinguished from S. merrillii by more robust plants, longer stem leaves (1.6–2.7 mm in length), much deep serration at upper leaf margins, and less weakly prorate laminal cells. In addition, as far as examining previously collected specimens in Asia, these two species have separate distributional areas (Fig. 3A). Plants collected in Halmahera Island, Indonesia (Vriese s.n.) have deeply concave leaves with scarcely developed proration on laminal cells. Plants collected in Luzon Island (Jacobs B415 and Jacobs B652) have quite slender, filamentous and pendulous secondary stems and they look similar to the pendulous form of Symphyodon recurvomarginatus. The latter, however, can be distinguished from S. merrillii by its totally smooth laminal cells and shorter costae (less than 1/5 of leaf length).

6-3. Symphyodon longispinosus H.Akiyama sp. nov. Figs. 3A & 11

Diagnosis: Similar to Symphyodon perrottetii in ovate leaves, distinct undulation in the upper part of leaves, lack of proration on laminal cells, and echinate cylindrical capsules, but differing in longer (to 200 µm in length) spines on exothecial cells. Type: CHINA. Yunnan Prov.: Tengchong Co., Western slopes of the southern portion of the Gaoligonshan. Irrawaddy River Basin, Yungeng Mountain National Cultural Park, along summit ridge trail from tram to mountain base. 23°22′49.6″N, 98°53′53.1″E, 2,070 m elev., Shevock 28681 (holotype CAS!) Etymology: This species was named after its long spines on capsule walls. Description: Plants yellowish green, more or less glossy. Primary stems pinnately branched, prostrate, attached to substrates. Secondary ascending stems irregularly and sparsely branching, to 3 cm long, densely and ± complanately foliate; central strands absent. Paraphyllia and pseudoparaphyllia absent. Dormant branch buds surrounded by widely triangular scaly leaves. Axillary hairs 100–200 µm long, with 2 (–3), pale brown, quadrate cells at base, and 3–7 hyaline, rectangular cells above. Asexual propagules absent. Stem and branch leaves of ascending stems not clearly differentiated, widely spreading when dry or wet, ovate or ovate-lanceolate, narrowed at base, acute, 1.6–2.2 mm long, ± concave, distinctly undulate above when dry or wet, narrowly decurrent at base; margins plane, sharply serrate above, serrulate below, almost entire at base; apical laminal cells short-rhomboidal to short- linear, 15–40 µm in length, smooth, plane; median and basal laminal cells linear, 50–80 µm in length, becoming shorter at both margins, smooth, plane; costae double, reaching 1/4–2/5 of leaf length; alar cells differentiated, quadrate to rectangular, slightly porose, moderately thick- walled. Calyptra in young stage cucullate and totally covering capsules, naked. Dioicous; perigonia and perichaetia on separate secondary stems. Perigonia with 7–10 antheridia, without paraphyses; leaves pale yellow, shortly ovate, 1 mm long, acute, concave, ecostate. Perichaetia with ca. 6–15 archegonia, with straight, uniseriate paraphyses; innermost leaves ovate, acuminate, 1.3 mm long, ecostate, lower laminal cells smooth, distinctly porose. Setae 6–11 mm long, reddish brown, densely papillose above, sparsely papillose or entire below. Operculum long-rostrate, 1.0–1.2 mm long, setose below, smooth above. Capsule erect,

80 Figure 11. Symphyodon longispinosus H.Akiyama. A–D: Leaves of ascending stems. E: Basal part of a leaf. F–G: Apices of stem leaves. H: Median laminal cells. I: Plants with sporophytes. J: Close-up of sporophytes: two young calyptrae, median part of a seta, and an echinate capsule. K: Lower part of outer surface of exostome teeth. L: Upper part of a capsule. M: Lower part of a capsule. N: Close-up of spines of a capsule. All from Shevock 28681 (holotype, CAS). Scale bars＝0.1 mm for A–C, D & E, 0.1 mm for E, 10 µm for F, H & K, 20 µm for G, 5 mm for I, 1 mm for J, 0.5 mm for L–M, 0.2 mm for N.

81 ovoid to short-cylindrical, 1.8–3.3 mm long, with 6–7 layers in transverse section, densely echinate (most spines 70–200 µm in length) ; exothecial cells long-rectangular, thick-walled; stomata present at base, few in number; annuli differentiated, of 6–7 rows with small, quadrate, very thick-walled cells. Peristome double; exostome teeth linear-lanceolate, whitish, to 300 µm tall, minutely papillose; endostome with low basal membrane, segments linear, without perforation, variable in length, 80–170 µm tall, cilia absent. Spores spherical or slightly ellipsoid, green, minutely papillose, 20–25 µm in diameter. Distinguishing features: 1) Irregularly branched ascending shoots with wide- spreading leaves in both dry and wet condition, 2) poor differentiation between stem and branch leaves, 3) ovate and acuminate leaves with distinct undulation above, 4) sharp serration of upper leaf margins, 5) smooth laminal cells, and 6) cylindrical capsules with long spines (to 200 µm in length). Habitat: Growing on trunks and branches of evergreen trees under filtered light at rather higher elevation (2,014–2,337 m elev.). Distribution: China (endemic to western part of Yunnan Prov.). Paratypes: CHINA. Yunnan Prov., Nanjin Co., Dali, Wuliangsha Mountain Natural Reserve, 24°45′42″N, 100°31′11″E, 2,337 m elev., Ma 12-2814 (CAS!, KUN; as Symphyodon perrottetii); Longyang Qu, Lujiang Xiang, Bawan Cun, east slope of Gaoligong Shan, below summit or ridge, near pass on road from Bawan to Tengchong, 35 km from Bawan, 24°55′32″N, 98°45′27″E, 2,210 m elev., Long 32323 (CAS!, E; as S. perrottetii); ibid., along new road from Tengchong to Lujiang, near Nankang rest house, 24°49′23″N, 98°46′55″E, 2,014 m elev., Long 32819 (CAS!, E; as S. perrottetii); Jin-Don Co., Wuliang Shan, Kunming Forest coll. 14 (KUN, L!; as S. perrottetii); ibid., Kunming University, Department of Biology 37 (KUN, L!; as S. perrottetii). Taxonomical notes: This new species is closely related to Symphyodon perrottetii in its ovate leaf shape with distinct undulation and sharp serrations at the upper margins, smooth laminal cells, and cylindrical capsules with dense spines. The latter, however, differs from the former in not so widely spreading stems, and shorter spines on the exothecial cells (less than 90 µm in length according to He & Snider 2000). All the specimens of Symphyodon longispinosus have been identified as S. perrottetii. Chinese specimens previously identified as such [e.g., those reported in Moss Flora of China (He & Lin 2002)] need to have their identity verified. According to Noguchi (1972), capsules of Symphyodon erinaceus are cylindrical bearing long spines and thus they look similar to those of the present new species. But S. erinaceus has a sparsely branching pattern and long-pendent plants (often reaching to 30 cm long) with deeply concave leaves, obtuse or obtusely pointed apices, and more or less prorate laminal cells, which do not suggest a close relationship between S. erinaceus and S. longispinosus. He & Snider (2000) wrote that spines on capsules of S. erinaceus do not exceed 85 µm in length. The ovate leaves and long spines (to 75 µm in length) on the exothecial cells of Symphyodon echinatus suggest its relationship to S. longispinosus. However, S. echinatus differs in regularly and densely pinnate–tripinnate branching pattern of ascending secondary stems, distinct differentiation between stem and branch leaves, short costae, and well-developed proration of laminal cells. In addition, the six specimens of S. longispinosus do not bear

82 filamentous asexual propagules, which have been reported for S. echinatus (He & Snider 2000). The long-echinate exothecial cells of Symphyodon longispinosus are similar to those of S. gollanioides (to 150 µm in length), which has been reported from northern Thailand, Sikkim and Japan (Noguchi 1972; He & Snider 2000; this report). They also share almost smooth laminal cells. However, S. gollanioides differs from S. longispinosus in its dense pinnate branching of ascending stems, shorter (1.1–1.6 mm), ovate to oblong-ovate, broadly acute to obtuse leaves, lack of marked undulation in the upper portions of leaves, and shorter (2.0– 2.2 mm long), ovoid capsules. For comparison with another species reported from Yunnan, China, S. yuennanensis Broth., see the note below.

6-4. Symphyodon yuennanensis Broth., Symb. Sin. 4: 90. 1929. Figs. 3A & 5L–O Type: CHINA. Yunnan, An Bäumen in der wtp. St. des birm. Mons. im Regenmischwald ober Schutsche am Djiou-djiang (Irrawadi-Oberlauf), 27°55′N, 2400–2800 m, Handel-Mazzetti 9473 (holotype H-BR!, isotypes H-BR!, S, W)

Distribution: Endemic to Guizhou and Yunnan in China. Other specimen examined: CHINA. Guizhou, Honghe Pref., Jinping Co., Zhemi Town Fenshuiling National Nature Reserve, Xilongshan Mts., Reserve Station from Sanchahe to the summit of Mt. Xilongshan (Phu Si Lung), 23°37.515′N, 102°47.376′E, 2,973 m, Wang et al. 20130826-96 (HSNU). Taxonomical notes: Symphyodon yuennanensis has been known only by single type specimen collected in northwestern Yunnan Province (He & Snider 2000), but an additional specimen has been confirmed from Guizhou, China (Fig. 3A). The species differs from S. longispinosus, newly desribed here based on the specimens collected from Yunnan, China, in slender plants with lightly appressed leaves, distinct proration of laminal cells, and weak serration at leaf margins (Fig. 5). According to He & Snider (2000), leaf apices of S. yuennanensis are bluntly acuminate and this feature was attributed as one of the distinguishing features of the species. Examination of type materials (holotype and isotype deposited in H-BR), however, reveals that they are not bluntly acuminate but acuminate (Fig. 5M–O) as given in the protologue (Brotherus 1929).

6-5. Symphyodon recurvomarginatus (Dixon & Sakurai) H.Akiyama, comb. nov. Figs. 3A–B & 12 ≡Glossadelphus recurvomarginatus Dixon & Sakurai, Bot. Mag. (Tokyo) 50: 620. 1936. Type: JAPAN. Kiushiu, Prov. Ohsumi, Mt. Takakuma, Doi s.n. in Herb. Sakurai 3549 (holotype MAK!) ＝Symphyodon perrottetii auct. non Mont., sensu Japanese bryologists (Ando & Seki 1962; Noguchi et al. 1991)

Description: Plants yellowish green, sometimes becoming slightly blackish, dull or faintly glossy. Primary stems shortly prostrate on substrates, densely branched. Secondary stems usually to 3 cm long, sometimes with pendulous and flagelliform tips, reaching 5–7 cm long. Paraphyllia and pseudoparaphyllia absent. Asexual propagules absent. Stem and branch

83 Figure 12. Symphyodon recurvomarginatus (Dixon & Sakurai) H.Akiyama. A–C: Plants. D: Close-up showing undulate leaves. E: Plant with a sporophyte. F: Capsule with low spines. G–L: Secondary stem leaves. M–N: Branch leaves. O–P: Apices of stem leaves. Q–R: Alar regions of stem leaves. S–U: Median laminal cells of stem leaves. A–B, I–J, R–S & V from Akiyama 22798 (HYO). C–D, K–N & U from Akiyama 22960 (HYO), E–F from Seki 29356 (HIRO), G–H, O, P & S from Doi s.n. (holotype of Glossadelphus recurvomarginatus, MAK). Scale bars＝5 mm for A–C, 2 mm for D & E, 100 µm for F– M, 20 µm for O–U. 84 leaves not so differentiated in size and shape. Branches short, to 1 cm long. Stem leaves ovate to lanceolate, 1.0–1.6 mm long, rarely reaching 2.0 mm long, more or less undulate above, acute to acuminate, serrulate at upper margins; upper laminal cells short-rhomboidal to linear, smooth; median laminal cells linear, smooth; costae double, 1/6–1/4 of leaf length; alar cells differentiated, quadrate. Branch leaves ovate-lanceolate to oblong, 0.8–1.4 mm long, more or less undulate above, serrulate to weakly serrate at upper margins; upper laminal cells short- rhomboidal to linear, smooth; median laminal cells linear, smooth; costae double, reaching 1/5–1/4 of leaf length; alar cells differentiated, few, quadrate. Calyptrae unknown. Dioicous. Female and male plants similar in size and branching. Perigonia at lower portion of secondary branches; with 5–6 antheridia; paraphyses absent. Perigonial leaves ovate, to 0.6 mm long. Perichaetia at the basal portion of a secondary branch. Post fertilized perichaetial leaves linear-lanceolate, to 1.5 mm long. Setae ca. 10 mm long, reddish brown, highly mammillose above, almost smooth below. Capsules erect, cylindrical, ca. 1.7–2.0 mm long, 0.5 mm wide, reddish brown; capsule walls with short, triangular spines, 30–75 µm in height (Fig. 12F). Peristome double, yellowish white. Exostome teeth linear-lanceolate, filamentous above, to 300 µm in height, smooth. Endostome segments linear, to 120 µm in length on low basal membrane; cilia absent. Other features unknown. Distinguishing features: 1) Soft plants with short branches not spreading horizontally, often pendulous from substrates, 2) stem leaves acute, serrulate to weakly serrate at upper leaf margins, 3) often undulate upper lamina, 4) totally smooth laminal cells in stem and branch leaves, and 4) low echination as in the case of Symphyodon merrillii and S. perrottetii. Habitat: Growing pendent from shrub twigs and branches in more or less windy places in warm temperate to subtropical montane forests with heavy annual rainfall. Distribution: Japan, Taiwan, and northern Vietnam. In Japan, Symphyodon recurvomarginatus is distributed in geographically isolated patches along the Pacific coast. The specimen collected at Kizu, Kaizan-cho, Mie Pref. (Takaki 28329) is the northern limit of this species. Representative specimens examined: JAPAN. Tokyo Met., Hachijojima Island, Mt. Nishiyama, Akiyama 22785 (HYO); Mt. Mihara, summit, Akiyama 22835 (HYO). Mikurajima Island, Sato-Borosawa, Higuchi 37392 (TNS). Mie Pref., Miyama-cho, upper reach of Choshi River, around Kizu, 200 m elev., Takaki 28320 (NICH). Tokushima Pref., Todoroki-Kujyuku- taki Falls, Torikaeshino-taki Fall, 410 m elev., Tateishi 32825 (HIRO); ibid., Nabewari Fall, 450 m elev., Nishimura 13226 (OKAM). Kumamoto Pref., Mt. Ichifusa, Akiyama 23592 (HYO). Miyazaki Pref., Tsuno-cho, Osuzu-keikoku Valley, Akiyama 22879 (HYO); Nichinan-shi, Sakatani, Inotani Valley, Akiyama 22960 (HYO); Shiiba-mura, Ohkochi, Nojinotaki Fall, 560 m elev., Tateishi 34625 (HIRO). Kagoshima Pref., Kimotsuki-gun, Kimotsuki-cho (Takayama-cho), Mt. Hutamata, 720 m elev., Seki 29356 (with a single sporophyte) (HIRO); Yakushima Island, Suzukawa Valley, 150–500 m elev., 7 Dec. 1964, Iwatsuki et al. s.n. (distributed as Musci Japonici no. 1043); ibid., Yakusugi-land, Akiyama 24774 (HYO); ibid., Mocchomu-dake, Ohgue & Hayashida 001 (HYO); ibid., Shiratani, Hayashida 383 (HYO); Amamiohshima Island, Mt. Yuwan-dake, around the summit, Akiyama 24400 (HYO); Tokunoshima Island, Mt. Inokawa-dake, around the summit, ca. 500 m elev., Akiyama 24490 (HYO). TAIWAN. Taichung City, Heping Dist., Dasyueshan National Forest

85 Recreation Area, along a trail between Police Station to Small Sacred Tree, 2,200 m elev., Akiyama 23285 (HYO). VIETNAM. Vinh Phuc Co., Tam Dao National Park, en route from the Lodge House (979 m elev.) to the summit of Mt. Tam Dao 1, 1,170 m elev., Akiyama 24138 (HYO); ibid., climbing approach to the Television Tower Mountain, 1,140 m elev., Akiyama 24157 (HYO). Cao Bang Co., Nguyen Binh Dist., Phia Oac-Phia Den Nature Reserve, 22°36′28.83″N, 105°52′14.76″E, 1,604 m elev., Zhu et al. 20131025-304 (HSNU). Taxonomical notes: Plants of Symphyodon recurvomarginatus are very soft in texture, and the irregularly and pinnately branched secondary stems often grow downward on the surface of tree trunks or hang from the twigs and branches of small shrubs. In addition, the tips of the drooping plants, including lateral branches do not roll up or spread horizontally as in the case of epiphytic mosses of similar plant appearance such as Neckera flexiramea Cardot. In the first report of the genus Symphyodon from the Japanese Islands, Ando & Seki (1962) treated Gollania recurvomarginatus Dixon & Sakurai as a synonym of Symphyodon perrottetii. Since then, He & Snider (2000) and other taxonomists have reported Symphyodon plants from Japan, China and Taiwan under the name of S. perrottetii. However, all plants of such reports are now revealed to be different from S. perrottetii s. str. as defined here. Of these, those plants from Chichibu region in Japan are recognized as S. copelandii, while those from Yunnan Prov. in China are S. longispinosus. Plants distributed in inland areas of Japan are treated as S. rheophilus and S. gollanioides. In addition, the plants previously recognized as S. perrottetii, distributed along the Pacific coast of Japan, such as Izu Islands, Wakayama, Kochi, Miyazaki, Kumamoto, and Kagoshima Pref. (including Yakushima Island and Amami Islands), correspond to Glossadelphus recurvomarginatus, which was described based on a specimen collected at Takakuma Mountains, Kagoshima Pref. As for gametophytic features of Symphyodon recurvomarginatus, Nishimura (2018) reported its morphological features in detail under the name of S. perrottetii using the specimen collected at Karei-tani valley, Kaiyo-cho, Tokushima Pref. (Tateishi 32713, HIRO, OKAY & TNS). So far, only two specimens with sporophytes have been found (Japan, Kagoshima Pref., Mt. Futamata, Seki 29356, HIRO; Japan, Kagoshima Pref., Yakushima Island, Kosugidani, Iwatsuki et al. 15245, NICH). As previously pointed out by Ando & Seki (1962), when Taxiphyllum taxirameum is growing on rocks in arid habitats, its plant morphology becomes similar to that of Symphyodon spp., such as in leaf shape, short and double costa, and weakly developed alar regions, and it sometimes becomes difficult to distinguish them by morphology alone. Plants collected at Mt. Amagi (Kagoshima Pref., Tokunoshima Island; Akiyama 24551, HYO) and at the Amano-Iwato, south of Mt. Asakuma (Mie Pref.; Iwatsuki et al. 10591, NICH) are good examples. The appearance of plants and leaf shapes of Symphyodon recurvomarginatus is quite variable and thus even the plants with the same DNA sequences have completely different leaf shapes. In terms of plant appearance, for example, the population at Hachijojima Island is particularly slender, while the plant at Inotani, Miyazaki is larger and flat. In addition, plants tend to become larger toward the southern part of their distributional areas, especially in Yakushima Island. In such case, it sometimes becomes difficult to distinguish this species from

86 S. merrillii, which is widely distributed in the Malesia region. However, S. recurvomarginatus is characterized by completely smooth laminal cells, whereas S. merrillii is distinguished by a tendency to produce proration, even if they are small. As mentioned above, the molecular phylogeny (Fig. 1) suggests that both of the two species are separate species. Symphyodon erraticus, reported from Bhutan, southwestern India and Sri Lanka, is similar to S. recurvomarginatus in its overall plant shape, narrower leaves and short costae. However, after examination of specimens collected from north India [Bretanderu 59 & 74 (H) and Hartees 2709 (H)], north Thailand [Akiyama & Printarakul 1465 (HYO)], Sri Lanka [Herzog 42 (H)] and north Vietnam [Zhu et al. 20131026-225A (HSNU)], we found that S. erraticus differs from S. recurvomarginatus in smaller plant size, densely bipinnate branching pattern, differentiation between stem and branch leaves, and amongst all, upper and median laminal cells with strong proration often with small spines.

Acknowledgments We thank Dr. K. Santanachote (Chiang Mai University) and Dr. N. Printarakul (Chiang Mai University) for their kind help in the field surveys and providing valuable plant samples, and Office of the National Research Council of Thailand (NRCT) for permission to do field research in Doi Inthanon National Park, Ms. Ida Haerida (LIPI-Cibinong) for her help in field work in Indonesia, Mr. James R. Shevock (California Academy of Sciences) for providing a number of fresh and valuable plants from Hawaii, Taiwan, and China (Yunnan) including Rheoshevockia fontana, Ms. T. T. Luong (University of Sciences, Vietnam) and Mr. D. N. Lam (Southern Institute of Ecology, Vietnam) for their kind help in fieldworks in North and South Vietnam, Dr. S.-H. Lin (Tunghai University, Taiwan), Dr. J.-D. Yang, and Dr. K.-Y. Yao (Endemic Species Research Institute, Taiwan; TAIE) for their help in fieldworks in Taiwan, Dr. S. He (Missouri Botanical Gardens) for his valuable advice, Ms. M. Saji for her kind guidance in field survey in Tokushima Pref., Dr. Y. Inoue (National Museum of Nature and Science), Dr. H. Kiguchi (Saitama Pref.), the late Mr. Y. Tateishi (Okayama Pref.), Dr. N. Nishimura (Okayama Pref.), Mr. N. Hayashida (Kagoshima Pref.), and Dr. M. Higuchi (National Museum of Nature and Science) for their kind offers of valuable specimens from various parts of Japan. We also thank to Dr. W. R. Buck (New York Botanical Garden) for linguistic corrections of the text. We are deeply debt to the curators of BM, BO, BORH, CAS, CMUB, E, FH, H, HIRO, HSNU, HYO, KUN, KYO, L, MAK, MO, NICH, NY, OKAM, OSA, PC, PE, SCCN, SNP, TAIE, TNM, TNS, and UC for their kind arrangement of valuable specimens. This study was financially supported by JSPS KAKENHI no. 24570108 to H. A. and no. 24255007 to Dr. M. Kanzaki (Kyoto University).

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Appendix 82 samples used in the analyses and their accession numbers (in order of rps4, trnL-F, rbcL, and ITS1&2). New accessions are indicated with an asterisk (*) before species names and with voucher information. Scientific names of registered data are shown for Symphyodon species. Symphyodontaceae: Chaetomitrium borneense AY306880, AY306714, —, HQ613413. C. dusenii AY306881, AY306715, —, HQ613414. *C. horridulum (VIETNAM. Bidup Nuiba N. P., Akiyama 23956, HYO) LC620860, LC620860, LC620177, LC620246. *C. orthorrhynchum (MALAYSIA. Sabah, Kinabalu Park, Akiyama 23432, HYO) LC620861, LC620861, LC620178, LC620247. C. orthorrhynchum HQ443821, HQ443859, —, HQ443722. Chaetomitriopsis glaucocarpa AY908603, HQ443858, —, HQ443721. Dimorphocladon borneense AY306898, AY306732, —, HQ443732. Phyllodon lingulatus AM990367, AM990367, —, FM161105. P. trucatulus AY908604, HQ443880, —, HQ443764. *Rheoshevockia fontana (CHINA. Yunnan Province, Shevock46478, CAS) MK257801, MK257805, LC620179, MK164022. R. fontana MK257802, MK257806, —, MK164023. *Symphyodon asper (TAIWAN. Miali, Akiyama 23149, HYO) LC620760, LC620829, LC620146, LC620216. *S. asper (TAIWAN. Miali, Akiyama 23231, HYO) LC620761, LC620830, LC620147, LC620217. *S. asper (TAIWAN. Miali, Akiyama 23254, HYO) LC620762, LC620831, LC620148, LC620218. *S. asper (JAPAN. Tokushima Pref., Todoroki Falls, Nishimura 13212, OKAM) LC620763, LC620832, LC620149, LC620219. *S. asper (JAPAN. Tokushima Pref., Todoroki Falls, Tateishi 33116, HIRO) LC620764, LC620833, LC620150, LC620220. *S. asper (JAPAN. Kochi Pref., Takeyashiki, Tateishi 35324, HIRO) LC620765, LC620834, LC620151, LC620221. *S. asper (JAPAN. Kumamoto Pref., Mt. Ichifusa, Akiyama 23594, HYO) LC620766, LC620835, LC620152, LC620222. *S. asper (JAPAN. Miyazaki Pref., Nichinan City, Akiyama 22959, HYO) LC620767, LC620836, LC620153, LC620223. *S. complanatus (VIETNAM. Bidup-NuiBa National Park, Lang Biang, Akiyama 25539, HYO) LC620784, LC620853, LC620170, LC620240. *S. copelandii (JAPAN. Saitama Pref., Ogano Town, Haccho Pass, Akiyama 22992, HYO) LC620728, LC620796, LC620115, LC620184. *S. copelandii (JAPAN. Saitama Pref., Ogano Town, Haccho Pass, Akiyama 22993, HYO) LC620729, LC620797, LC620116, LC620185. *S.

90 copelandii (JAPAN. Saitama Pref., Ogano Town, Haccho Pass, Akiyama 22994, HYO) LC620730, LC620798, LC620117, LC620186. *S. copelandii (JAPAN. Saitama Pref., Otaki Village, Otaki, Akiyama 23014, HYO) LC620731, LC620799, LC620118, LC620187. *S. copelandii (JAPAN. Saitama Pref., Otaki Village, Otaki, Akiyama 23016, HYO) LC620732, LC620800, LC620119, LC620188. *S. chrysobasilatus (U.S.A. Hawaii Islands, Kauai Island, Shevock et al. 46302, CAS) LC620787, LC620857, LC620174, LC620244. *S. erraticus (THAILAND. Doi Inthanon, Akiyama & Printarakul 1465, HYO) LC620794, LC620794, LC620113, LC620182. *S. erinaceus (THAILAND. Doi Inthanon, Akiyama & Printarakul 1460, HYO) LC620801, LC620801, LC620120, LC620189. *S. erinaceus (VIETNAM. Bidup-NuiBa National Park, Lang Biang, Akiyama 25540, HYO) LC620855, LC620855, LC620172, LC620242. *S. gollanioides (JAPAN. Mie Pref., Miyanotani, Akiyama 21857, HYO) LC620795, LC620795, LC620114, LC620183. S. imbricatifolius AY306999, AY306833, —, HQ443774. *S. imbricatifolius (COLOMBIA. Antioquia, Cerro de la Vieja, Churchill & Betancur 18734, MO) LC620809, LC620809, LC620128, LC620197. S. imbricatifolius MK257803, MK257807, —, MK164024. *S. leiocarpus (THAILAND. Doi Inthanon, Akiyama & Pritarakul 1472, HYO) LC620810, LC620810, LC620129, LC620198. *S leiocarpus (THAILAND. Doi Inthanon, Akiyama et al. 1275, HYO) LC620811, LC620811, LC620130, LC620199. *S. longispinosus (CHINA. Yunnan Prov., Long 32323, CAS) LC620812, LC620812, LC620131, LC620200. *S. longispinosus (CHINA. Yunnan Prov., Long 32819, CAS) LC620813, LC620813, LC620132, LC620201. *S. longispinosus (CHINA. Yunnan Prov., Ma 12-2814, CAS) LC620814, LC620814, LC620133, LC620202. *S. longispinosus (CHINA. Yunnan Prov., Shevock 28681, CAS) LC620815, LC620815, LC620134, —. *S. merrillii (TAIWAN. Taoyuan Co., Shevock 30525, HYO) LC620817, LC620817, LC620136, LC620204. *S. merrillii (PHILIPPINES. Mindanao, Shevock 44714, HYO) LC620818, LC620818, LC620137, LC620205. *S. merrillii (PHILIPPINES. Mindanao, Shevock 44824, HYO) LC620819, LC620819, LC620138, LC620206. *S. merrillii (INDONESIA. Lombok Island, Touw & Snoek 23213, L) LC620786, LC620856, LC620173, LC620243. *S. ovalifolius (MALAYSIA. Sabah, Kinabalu Park, Akiyama & Suleiman 23433, HYO) LC620803, LC620803, LC620122, LC620191. *S. ovalifolus (MALAYSIA. Sabah, Kinabalu Park, Akiyama & Suleiman 23463, HYO) LC620804, LC620804, LC620123, LC620192. *S. ovalifolius (MALAYSIA. Sabah, Kinabalu Park, Akiyama & Suleiman 23464, HYO) LC620805, LC620805, LC620124, LC620193. *S. ovalifolius (MALAYSIA. Sabah, Kinabalu Park, Akiyama & Suleiman 23466, HYO) LC620806, LC620806, LC620125, LC620194. *S ovalifolius (INDONESIA. Java, Gunung Gedeh, Akiyama & Haerida 24856, HYO) LC620807, LC620807, LC620126, LC620195. *S. ovalifolius (INDONESIA. Java, Gunung Gedeh, Akiyama & Haerida 24858, HYO) LC620808, LC620808, LC620127, LC620195. *S. pygmaeus (U.S.A. Hawaii Islands, Oahu Island, Shevock et al. 44534, CAS) LC620858, LC620858, LC620175, —. *S. pygmaeus (THAILAND. Doi Suthep, Printarakul 190420, HYO) LC620859, LC620859, LC620176, LC620245. *S. recurvomarginatus (JAPAN. Tokyo Metro., Hachijojima Island, Mt. Nishiyama, Akiyama 22785, HYO) LC620837, LC620837, LC620154, LC620224. *S. recurvomarginatus (JAPAN. Tokyo Metro., Hachijojima Island, Mt. Mihara, Akiyama 22835, HYO) LC620838, LC620838, LC620155, LC620225. *S. recurvomarginatus (JAPAN. Miyazaki Pref., Osuzu Valley,

91 Akiyama 22879, HYO) LC620839, LC620839, LC620156, LC620226. *S. recurvomarginatus (JAPAN. Miyazaki Pref., Nichinan City, Akiyama 22960, HYO) LC620840, LC620840, LC620157, LC620227. *S. recurvomarginatus (JAPAN. Kumamoto Pref., Mt. Ichifusa, Akiyama 23592, HYO) LC620841, LC620841, LC620158, LC620228. *S. recurvomarginatus (VIETNAM. Tam Dao National Park, Akiyama 24138, HYO) LC620842, LC620842, LC620159, LC620229. *S. recurvomarginatus (VIETNAM. Tam Dao National Park, Akiyama 24157, HYO) LC620843, LC620843, LC620160, LC620230. *S. recurvomarginatus (JAPAN. Kagoshima Pref., Amami Island, Akiyama 24400, HYO) LC620844, LC620844, LC620161, LC620231. *S. recurvomarginatus (JAPAN. Kagoshima Pref., Tokunoshima Island, Akiyama 24490, HYO) LC620845, LC620845, LC620162, LC620232. *S. recurvomarginatus (JAPAN. Tokushima Pref., Todoroki Falls, Nishimura 13226, OKAM) LC620846, LC620846, LC620163, LC620233. *S. recurvomarginatus (JAPAN. Miyazaki Pref., Shiiba Village, Tateishi 34625, HIRO) LC620847, LC620847, LC620164, LC620234. *S. recurvomarginatus (JAPAN. Kagoshima Pref., Yakushima Island, Mt. Mocchomu, Ohgue & Hayashida 001, HYO) LC620848, LC620848, LC620165, LC620235. *S. recurvomarginatus (JAPAN. Kagoshima Pref., Yakushima Island, Shiratani Valley, Hayashida 383, HYO) LC620849, LC620849, LC620166, LC620236. *S. rheophilus (JAPAN. Mie Pref., Miyanotani, Akiyama 21855, HYO) LC620820, LC620820, LC620139, LC620207. *S. rheophilus (JAPAN. Mie Pref., Miyanotani, Akiyama 21858, HYO) LC620821, LC620821, LC620140, LC620208. *S. rheophilus (JAPAN. Mie Pref., Miyanotani, Akiyama 21861, HYO) LC620822, LC620822, LC620141, LC620209. *S. rheophilus (JAPAN. Mie Pref., Miyanotani, Akiyama 21863, HYO) LC620823, LC620823, —, LC620210. *S. rheophilus (JAPAN. Tokushima Pref., Todoroki Falls, Tateishi 33138, HIRO) LC620824, LC620824, LC620142, LC620211. *S. rheophilus (JAPAN. Tokushima Pref., Todoroki Falls, Nishimura 13225, OKAM) LC620825, LC620825, LC620143, LC620212. *S. rheophilus (JAPAN. Miyazaki Pref., Osuzu Valley, Akiyama 22892, HYO) LC620826, LC620826, LC620144, LC620213. *S. rheophilus (JAPAN. Miyazaki Pref., Osuzu Valley, Akiyama 22870, HYO) LC620827, LC620827, —, LC620214. *S. rheophilus (JAPAN. Miyazaki Pref., Osuzu Valley, Akiyama 22875, HYO) LC620828, LC620828, LC620145, LC620215. *S. scaber (VIETNAM. Bidup-NuiBa National Park, Lang Biang, Akiyama 25542, HYO), —, LC620816, LC620135, LC620203. *S. scaber (VIETNAM. Bidup-NuiBa National Park, Lang Biang, Akiyama 25543, HYO), LC620854, LC620854, LC620171, LC620241. *S. scabrisetus (VIETNAM. Bidup NuiBa National Park, Akiyama 23977, HYO) LC620850, LC620850, LC620167, LC620237. *S. scabrisetus (VIETNAM. Bidup-NuiBa National Park, Akiyama 24005, HYO) LC620851, LC620851, LC620168, LC620238. *S. scabrisetus (VIETNAM. Bidup-NuiBa National Park, Akiyama 24064, HYO) LC620852, LC620852, LC620169, LC620239. *S. scabrisetus (THAILAND. Doi Inthanon, Akiyama & Printarakul 1493, HYO) LC620734, LC620802, LC620121, LC620190. Outgroup: Homaliodendron scalpelifolium HQ607402, —, AB094788, FM161135. *Filibryum ogatae (JAPAN. Saitama Pref., Otaki Village, Otaki, Akiyama 23015, HYO) LC620862, LC620862, LC620180, LC620248. *Taxiphyllum taxirameum (JAPAN. Kagoshima Pref., Tokunoshima Island, Mt. Inokawa, Akiyama 24505, HYO) LC620863, LC620863, LC620181, LC620249.

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