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Orthodontopsis, a New Genus of Bryaceae (Musci) from Southern Siberia, Ussr

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Orthodontopsis, a New Genus of Bryaceae (Musci) from Southern Siberia, Ussr Journ. Hattori Bot. Lab. No. 71: 165- 173 (Jan. 1992) ORTHODONTOPSIS, A NEW GENUS OF BRYACEAE (MUSCI) FROM SOUTHERN SIBERIA, USSR 1 2 MICHAELS. IGNATOV AND BENITO C. TAN Among the many bryophyte collections made recently by the senior author from the Altai mountain in southern Siberia, USSR, is an Orthodontium-like moss which proves to be new to science. The specimens were collected from a rotten log with Dicranum fragi/ifo/ium Lindb., Lepidozia reptans (Hedw.) Oum., Lophozia ventricosa (Dicks.) Oum. in subalpine Pin us sibirica forest on a south-facing slope at an elevation of 2100m in close proximity to the permafrost area. It grows also on soil under the roots of fallen trunks. The locality is Ayulyuyuzyuk Creek, Karakem River Basin in Central Altai (50 30'N - 89 lO'E). Later, Dr. L. V. Bardunov of Irkutsk Herbarium showed us his collection of the same species from Western Sayan Mt. adjacent to the Altai range, also from rotten wood at an elevation of 1700 m elevation in a Pin us sibirica and Larix sibirica forest. Both specimens have strong resemblance to species of Orthodontium, especially when dry. The gametophytes are erect, with loosely tufted plant habit, lanceolate-linear leaves, and the sporophytes are terminal in position. However, under the microscope, the leaf areolation is like a Bryum with rhomboidal to rectangular cells, except at the well developed leaf border. The capsule which is erect and also Orthodontium-like in shape and appearance has only a highly reduced endostome consisting of short and slender processes that are deeply inserted at the mouth and without a basal membrane (Plates 3 & 4). No remnants of an exostome is observed. Using the worldwide keys and descriptions contained in Brotherus ( 1924) and other regional floras or monographs, such as Fleischer ( 1902- 1904), Sainsbury (1955), Nyholm (1958), Ochi (1959), Lawton (1971), Gangulee (1974), Smith (1978), Crum and Anderson (1981), Koponen and Norris (1985), and Noguchi ( 1988), we are led to Orthodontium Schwaegr. (including Stableria Lindb.) or Mieli­ chhoferia Nees & Hornsch. The genus Orthodontium Schwaegr. has been treated in a worldwide monograph by Meijer (1952), who accepts eight species distributed chiefly in temperate oceanic and tropiCal mountainous climates, especially in the southern hemisphere. It is quite surprising therefore to find an Orthodontium-like moss in southern Siberia where there is a strong continental climate. Our Siberian moss is similar to species of Orthodontium in many respects: epixylic habitat, overall plant size, the branching pattern, the presence of fragile or deciduous branches, agreement in leaf outline, a spreading foliation with leaves often 3- 4 times twisted or flexuose, the costal anatomy, the autoicous condition and the papillose 1 Main Botanical Garden, Botanicheskaya 4, Moscow, USSR 127276. 2 Farlow Herbarium, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts, USA 02138. 166 Journ. Hattori Bot. Lab. No. 71 I 9 9 2 spores. All of these are strong indications that the new taxon may have evolved in isolation from an ancestral Orthodontium . However, all species of this genus, including the four presently known in northern hemisphere, namely, 0. lineare Schwaegr., 0. pellucens (Hook.) B. S. G ., 0. gracile Schwaegr. ex Bruch & Schimp. and 0. infractum Dozy & Molk., differ significantly from the Siberian moss in having ( 1) stems with a central strand, (2) unbordered leaves, (3) mostly elongate to linear laminal cells, and ( 4) a double peristome, albeit in various states of reduction. Obviously, to place the specimens in Orthodontium would require a basic alteration of its generic concept. The well differentiated and often bistratose leaf borders of the Siberian moss need additional comment. Consisting of 2 to 3 rows of elongate-linear cells, the bistratose borders occur on nearly all leaf margins except at the apex and proximal base. Within the subfamily Orthodontioideae, this is undoubtedly a disparate, autapomorphic char­ acter. We also compared this intriguing moss with the northern hemisphere species of Mielichhoferia and Asiatic members of Schizymenium Harv. ex Hook., the two genera of subfamily Mielichhoferioideae defined principally by the presence of a single peristome either representing a reduced exostome or endostome (see also Shaw 1985). Among those studied, the peristome structure of M. macrocarpa (Hook. ex Drumm.) Bruch & Schimp. (see Shaw & Crum 1984), M. himalayana Mitt. and S. javanicum (Broth. ex Fleisch.) Shaw come closest in their overall structural morphology. Nonetheless, the highly reduced, single structure observed in Mielichhoferia is the exostome (Shaw 1985) and not the endostome (cf. Shaw & Rohrer 1984) as is the case of Siberian moss. In Schizymenium javanicum, the single and endostomial peristome has a low basal membrane with remnants of cilium formation, and is quite different from that seen in our undescribed moss. It would appear that in bryaceous genera reduction of peristome into a single structure, either exostomial or endostomial, has evolved independently several times in separate lines of evolution. To force the Siberian moss into one of these two genera on the basis of a single peristome would create paraphyletic genera (see Fig. 1). In addition, species of Mielichhoferia and Schizymenium differ fundamentally from the Siberian moss in having ( 1) a caespitosely branching habit, (2) uniformly ovate­ lanceolate (rarely oblong to narrowly lanceolate) leaves, (3) laterally or basally located perichaetia, (4) thick, pyriformis capsules with well developed annuli, and (5) smooth to slightly papillose spores. In contrast, the moss under study has terminal inflores­ cences, although a few may grow on short lateral branch; and moreover, many PLATE l. Orthodontopsis bardunovii (based on holotype). A. Plant habit in wet condition; Al. The same plant with detached branches; A' : diagramatic scheme of the same plant showing position of gametangia; A2: autoicous branch with a secondary branch bearing antheridia in leaf axil (A2'). B. Upper portion of stem with leaves mostly removed to show the terminal and lateral archegonia. C. Branch with an enlarged, fertilized archegoniun. D. Leaves in dry condition. E. Stem leaves. F. Stem cross-section. G. Axillary hairs. H. Rhizoids. M. S. IGNATOV & B. C. TAN: Orthodontopsis, a new genus of Bryaceae 167 E E c N c:i ~ ~ \H G 168 Journ. Hattori Bot. Lab. No. 71 I 9 9 2 ~~ ~ c M. S. IGNATOV & B. C. TAN: Orthodontopsis, a new genus of Bryaceae 169 3 archegonia are seen scattered along the stem without subtending perichaetial leaves • Furthermore, its capsular wall is thin, the annulus is not apparent, and the spores are clearly papillose (see Plate 4c). The exothecial cells of Mielichhoferia and Schizym­ enium also are rounded, polygonal, quadrate to short rectangular in shapes, and with thick walls, whereas our problematic moss, like species of Orthodontium, has exothecial cells that are rectangular and oblong in shapes and have thin walls. Additionally, species of Mielichhoferia are epilithic and none is primarily epixylic on rotten wood, like this Siberian moss. We sent a specimen, labelled "Orthodontium sp.", to Dr. H. Ochi for his taxonomic opinion. His reply (pers. comm., 1991) was, "I have no name. I also tried Mielich­ hoferia, but the species of this genus all have a different leaf shape, and their leaves have long cells and no borders.... " We believe that our moss specimen has sufficiently distinctive combinations of characters and that it deserves a generic recognition setting it apart from Orthodontium sensu Meijer ( 1952) and Mielichhoferia and Schizymenium sensu Shaw & Crum ( 1982, 1984) and Shaw (1985). A less satisfactory decision would be to describe it as a new subgenus in Orthodontium or Bryum. But to do so would only make the two contending genera polyphyletic and blur the traditional definition of genera in the family Bryaceae. We propose, therefore, a new monotypic genus to accomodate this unique Siberian moss pending a phylogenetic revision of bryaceous genera on a worldwide basis. Orthodontopsis bardunovii lgnatov & Tan, gen. et spec. nova. Plates 1- 4. Caulis valde ramificans fasciculo centrali nullo. Folia basalia diminuta, versus apicem accrescentia, superiora linearia, breviter acuminata, margine ad apicem parce crenata, limbo bistratoso e cellulis bi­ triseriatis formato. Costa infra apicem folii evanescens. Cellulae laminorum rhomboideae, 1: 2-4. Autoicous. Perichaetia terminalia, rarius lateralia aeque ac archaegonia nuda lateraliter disposita. Capsula subcylindrica, pallida, membranacea. Operculum humiliter conucum obtusum. Annulus et peristomum extemum desunt. Peristomii intemi 16, perforati vet integri, membrana peristomii nulla. Sporae papillosae. Plants in tufted clusters, forming mat or growing intermixed among other bryophytes. Stems 3- 5 mm long, 0.2--0.3 mm wide, without a central strand and a differentiated cortical layer, branched mostly at base, with fragile branches equal in length to the main stem or 2-3 times shorter. Rhizoids well developed, mostly colorless, with a few colored ones, profusely branched, ea. I0-15µm wide, at times only 3-4µm wide, thin-walled, smooth, rarely verrucose. Leaves dull green, spreading when moist, flexuose to strongly spiral-twisted when dry, PLATE 2. Orthodontopsis bardunovii (based on holotype). A- J. Leaf cross-sections; K, L: leaves; Kl, K2, K3, LI & L2: leaf cells (position on the leaf indicated by arrows); M: medial lamina! cells. 3 Though
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