ON TWO EAST ASIAN SPECIES of BRACHYTHECIUM (BRACHYTHECIACEAE, Musei)
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J. Hallori Bot. Lab. No. 100: 191- 199 (Aug. 2006) ON TWO EAST ASIAN SPECIES OF BRACHYTHECIUM (BRACHYTHECIACEAE, MUSeI) MICHAEL S. IGNATov\ IRINA A. MILYUTINA2 AND SANNA HUTTUNEN3 ABSTRACT. Brachythecium auriculatum A. Jaeger was synonymized with Palamocladium leskeoides by Hoffman (1998). The present study shows their independence. Brachythecium complanatum Broth. is reported from Japan for the first time; this species was confused before with the super ficially similar B. garovaglioides (= B. wichurae). Descriptions and illustrations of both species are provided. Japanese species of Brachytheciaceae were carefully revised by Takaki (1955a,b; 1956), and later the taxonomic concepts of Takaki were followed by Noguchi (1991) with relatively minor changes. In the course of study of Brachythecium species of Asia we found new data which are important for understanding two of the species occurring in Japan. Brachythecium auriculatum A. Jaeger, Ber. Thiitigk. St. Gallischen Naturwiss. Ges. 1876- 1877: 340. l878.-Hypnum auriculatum Lindb., Acta Soc. Sci. Fenn. 10: 250. 1872, nom. illeg. - Camptothecium auriculatum (A. Jaeger) Broth., Nat. Pflanzenfam. (ed. 2) 11 : 353. 1925. Fig. l. Holotype: Brachythecium auriculatum LINDB. c. fro Sachalien, Dui, inter Dicranum majus, Maji 1861 Glehn (H-SOL 1157001!). Isotypes: BM!, LE! Plants medium-sized to rather robust, yellowish-green, relatively rigid. Stem regularly pinnate-branched, branches 10-15 mm long; foliage rather dense, subjulaceous. Axillary hairs 2- 3 celled, 45- 65 X9- 1O flm, upper cell up to 40 flm long. Pseudoparaphyllia triangu lar, acute, serrate. Stem leaves erect to slightly falcate, 2.0-2.4XO.9- 1.1 mm, lanceolate, gradually acuminate, widest in the lower 1110 of the leaf length, auriculate, deeply plicate; costa reaching 0.35-0.65 leaf length, 30- 70 flm wide at base, ending without spine; margin serrate throughout, especially in acumen and near auricules or, in some leaves, serrulate to subentire. Mid-leaf cells 35-65( -80)X6-9 flm, thick-walled, marginal cells relatively short, 35-40 flm long and quite even along most of the leaf margin; basal cells slightly wider, in the leaf corners with numerous shortly ovate and subquadrate cells, forming con spicuous auricules, extending beyond general leaf outline above decurrencies. Branch leaves more or less falcate, shorter and relatively narrower than stem leaves. Dioicous. Perichaetial leaves gradually or abruptly tapered to a narrow acumen, margin in transition to acumen often with multicellular teeth; costa short and weak. Paraphyses more or less I Main Botanical Garden of Russian Academy of Sciences, Botanicheskaya 4, Moscow 127276, Russia. E-mail: [email protected] 2 A. N. Belozersky' Institute of Physicochemical Biology, Moscow State University, Moscow 119992, Russia. E-mail: [email protected] 3 Botanical Museum and Division of Systematic Biology, P.O. Box 7 and 65, FIN-OOOI4 Universi ty of Helsinki, Finland. E-mail: [email protected] 192 J. Hattori Bot. Lab. No. lOO 2 006 H L 100 flm Fig. 1. Brachythecium auriculatum A. Jaeger (from holotype): A, B, E - habit; C - perichaetium; D - capsule (peristome broken) and operculum; F, G - median laminal cells; H, I - stem leaves; J-L - branch leaves; M, N - auriculate leaf base; 0 - basal laminal cells; P - cells at margin of perichaetial leaf in the transition from sheating base to acumen; Q-T - perichaetialleaves. Scale bars: I cm for E; 2mm for A-D; 1 mm for H-L and Q-T; 0.5 mm for M-N; 100 Jim for F-G, O-P. M. S. IGNATOV: On two East Asian species of Brachythecium 193 abundant, conspicuously exserted from perichaetium. Seta 10- 13 mm long, strongly rough ened throughout. Capsule inclined to horizontal, curved, ca. 1.5 mm long (without opercu lum). Operculum conic. Peristome complete, endostome basal membrane with cilia equal in length to segments. Calyptra when young with few hairs. Brachythecium auriculatum was first described from Sakhalin (Sahalien, Sachalien) Island by Lindberg (1872), as 'Hypnum (Brachythecium) auriculatum' Lindb. nom. illeg., and subsequently validated by A. Jaeger (Jaeger & Sauerbeck, 1878). The species has been accepted in the genus Camptothecium (Brotherus, 1925; Takaki, 1955a; Wang & Hu, 2005) or Brachythecium (Noguchi, 1991; Ignatov & Huttunen, 2002). Hoffmann (1997, 1998) in her world revision of the genera Homalothecium and Palamocladium synonymized Brachythecium auriculatum with the pantropical Palamocladium leskeoides. The lectotype of B. auriculatum was selected by her among the specimens from BM. We disagree with both synonymization and lectotypification made by Hoffmann (1998). The difference between these two species is apparent already from the description: Brachythecium auriculatum has rough seta, whereas the smooth seta is one of the diagnos tic characters of the genus Palamocladium. Specimens in BM lack sporophytes. However, sporophytes are present in specimens in the herbarium of S.O.Lindberg in Helsinki Univer sity (H-SOL) and in LE. It seems that Hoffmann' selection was done because she failed to find this specimen in Helsinki Herbarium, which is understandable because it is kept in a folder labelled "Hypnum latifolium". Specimens in H-SOL, BM and LE have identical la bels and the plants are also very similar. The specimen in H-SOL must, therefore, be con sidered as the holotype. Palamocladium leskeoides is similar to B. auriculatum in having auriculate leaf bases, but it differs with the costa reaching 0.75-0.95 of the leaf length (0.35-0.65 in B. auricula tum) and in having a smooth (not rough) seta. Brachythecium auriculatum is a rare species in the Russian Far East and only a few additional collections were seen, not allowing a full understanding of the variation of this species. The main character is the auriculate leaf base. In the holotype and the few speci mens listed below this character is fairly obvious and also the angular region of leaf is strongly undulate, forming folds when mounted on a slide (Fig. IH-O). Very typical are plants from Hokkaido illustrated by Takaki (1955a, Fig. 6: 1-5). However, there are speci mens that have a Brachythecium salebrosum-like appearance with very deeply plicate leaves and quite a large group of small alar cells, and where the angular region is almost flat. Such plants were collected, for example, in Kamchatka. The exact identity remains equivocal until more comprehensive material becomes available. Thus, records from re gions other than Sakhalin, Kurils and Hokkaido may also belong to other species. Thus, further studies are needed to confirm the occurrence of B. auriculatum in Honshu, China and the continental part of the Russian Far East. Brachythecium auriculatum belongs to basal group of Brachythecium that character ized by a rough seta (Appendix). Specimens examined: SAKHA Ll N ISLAND . Makarovo, colI. A. Bukhteeva 1956 (LE); Okhotskoe, colI. V. Ya. Ardeeva I.IX.1966 (IRK, MHA). KUR IL ISLA NS . Iturup Island, between Kurilsk and Pio ner, colI. L. Bardunov & Presman, 10.IXl980 (IRK, LE); Kunashir Island, colI. Malyshev 194 J. Hattori Bot. Lab. No. lOO 2 0 0 6 26.IX.1964 (IRK, MHA); Kunashir Island, mys Sukacheva, colI. V. Cherdantseva 12.VIII. 1978 (VLAD, LE). Brachythecium complanatum Broth., Rev. Bryol. n. s. 2: I!. 1929. Fig. 2 B, E, F, H. Holotype: Amur, distr. Zeisko-Bureinsk, Mikhailofskoje, 25 .VI.l910 Korotki (H-BR!). Isotype LE! Plants robust, light-green to whitish-yellow. Stem ± regularly or quite remotely pinnate-branched, branches to 10- 13 mm long; foliage rather dense, subjulaceous to subcomplanate. Axillary hairs 2- 3 celled, 35- 55 X 7 -I 0 j1m, upper cell up to 25 j1m long. Pseudoparaphyllia triangular, acute. Stem leaves erect to erecto-patent, straight or some times slightly falcate, 2.2- 3.0± 1.0- 1.3 mm, ovate-lanceolate, gradually to moderately abruptly acuminate, widest at 1/10-117 of leaf length, concave, wrinkly-plicate; costa reaching 0.4- 0.7 leaf length, 30-70(-90)j1m wide at base, ending without spine; margin serrate above, serrulate to subentire towards the base. Mid-leaf cells 55- 90(- 110) X 6-9 j1m , basal cells wider, in leaf corners not clearly delimited or forming an indistinct square group of short-rectangular to subquadrate cells. Branch leaves smaller; margin more serrate. Dioicous. Perichaetia and perigonia borne on stems. Perichaetialleaves abruptly ta pered to a narrow and distally filiform acumen; costa short and weak. Paraphyses numer ous, conspicuously exserted from perichaetium. Seta 13 mm long, strongly roughened. Capsule inclined, 1.8 mm long without operculum. Calyptra with sparse (2-4) hairs. Spores 20- 22j1m. This species was described from the Amur region of the Russian Far East, but re mained poorly known for a long time. When preparing the "Check-list of mosses of the former USSR", Ignatov and Afonina (1992) referred some specimens from Asian Russia to B. wichurae, a superficially similar taxon. Using the keys of Takaki (1955b) one might eas ily identify sterile B. complanatum as B. wichurae. Ignatov (1998), however found that all specimens identified as B. wichurae from Siberia and the Russian Far East belong in fact to B. complanatum. Thus, B. wichurae has to be excluded from the Russian bryoflora. Recent ly the same misidentification was found among the specimens issued as 'Brachythecium wichurae' in the exsiccate series "Musci Japonici". Most specimens called 'B. wichurae' (# 92, 560, 664, 1107) are correctly identified, but have to be called B. garovaglioides Broth. et Par., which is an earlier name for B. wichurae (cf. Ignatov & Koponen, 1996; Wang & Hu, 1998). However # 1004 has rough setae and agrees with B. complanatum in all other details (Fig. 2). The difference between B. complanatum and B. garovaglioides are summa rized in Table I. However, there are a lot of 'rather' and 'mostly' which indicates that scanty collections and undeveloped plants would be difficult if not impossible to identify by morphology alone.