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19 Tropical Bryology 3: 19-28, 1990

Gradsteinia andicola, a remarkable aquatic from South America

Ryszard Ochyra Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31- 512 Kraków, Poland

(Studies on Colombian Cryptogams XLI)

Abstract. A new moss genus and species, Gradsteinia andicola, is described from the northern Andes of Colombia. It is an aquatic moss known sterile and characterized by 1) oblong or oblong-ovate, concave, cucullate and recurved-apiculate leaves with a very strong and variable costa that is basically single but commonly repeatedly branched and spurred from the base, giving the leaves a polycostate appearance; 2) thick-walled, porose and irregularly uni- to multistratose lamina cells; 3) bicellular axillary hairs; 4) the presence of incomplete limbidia; 5) the absence of paraphyllia, pseudoparaphyllia, central strand and alar cells. Until the sporophyte of Gradsteinia becomes known, this very distinct genus is tentatively placed in the family Donrichardsiaceae, based primarily upon the presence of variously multistratose leaf laminae and leaf areolation.

In the course of my examination of aquatic pleurocarpous for my treatment of the for Flora Neotropica, Gradsteinia andicola Ochyra, gen. & I was surprised by a specimen collected at spec. nov. (Figs. 1-3) altimontane elevations in Colombia that was unlike any of the known genera of Plantae aquaticae, sat robustae, fragiles, aquatic pleurocarps. I feel confident in flavo-virides vel fuscae, nitentes. Caulis proposing a new species and genus for its elongatus, erectus, rigidus, simplex vel inception, although the material available parce irregulariter ramificatus, 13 vel is sterile and sexual organs are not pluria cm longus, pilis axillaribus observed. For reasons discussed below I bicellularibus, paraphylliis et tentatively place Gradsteinia in the small pseudoparaphylliis nullis, in sectione family Donrichardsiaceae, close to transversa ovalis vel ellipticus, e cellulis Donrichardsia Crum & Anderson, until externis 2-3(-4)-stratosis, minoribus, more material and sporophytes become fuscis, parietibus valde incrassatis, internis available. The generic name honours my 4-5-stratosis, magnis, hyalinis, parietibus friend S. Rob Gradstein, the Deputy sat crassis, fasciculo centrali nullo. Folia Director of Cryptogams in the Flora Neo- caulinaria et ramealia similia, sat remote tropica project, and is a tribute to his disposita, siccitate erecto-patentia, contribution to world and in particular curvata vel leviter convoluta, madefacta neotropical bryology. laxe imbricata, carinato-concava, 20 incurvata, non decurrentia, (1.1-)1.7-2.0 central strand absent; paraphyllia absent; mm longa, (0.4-)0.6-0.8 mm lata, oblonga buds scattered along stem or in leaf axils, vel oblongo-ovata, apice rotundata, consisting of a small branch primordium cucullata minute recurvato-apiculata, covered with reduced leaves; pseudopa- marginibus erectis, integris vel minutissi- raphyllia none; axillary hairs infrequent, me distanter parce serrullata in parte bicellular, brown, short. Stem and branch apicali, cellulis laminae irregulariter leaves similar, shrivelled, curved and rolled fluctuante uni- vel multistratosis, pellucidis when dry, rather distant, irregularly erect- vel obscuris, elongato-hexagonalibus vel spreading to patent, loosely imbricate, oblongo-rhomboidalibus, supra costas navicular and distinctly incurved when lineari-flexuosis, 50-90 µm longis, 7.5- wet giving the stem and branches a 10.5 µm latis, parietibus crassis, porosis, catenulate appearance, (1.1-)1.7-2.0 mm in parte folii infima fuscescenti- long, (0.4-)0.6-0.8 mm wide, non decur- lutescentibus, rhomboidalibus vel breviter rent, usually slightly narrowed at base, rectangularibus, 10-15 µm latis, 15-25 oblong or oblong-ovate, concave, rounded µm longis, parietibus valde incrassatis and cucullate at the apex with most leaves porosisque, cellulis folii angularibus aliis in the upper part of shoots recurved- conformibus, costis validissimis, valde apiculate, mostly eroded and fimbriate in variabilibus, callosis, dorso alatis, cristatis, the older part of the stem and branches; 1/3-4/5 basi folii occupantibus, bifurcis margins entire or distantly minutely vel pluripartitis, basi coalitis, ramis dein serrulate above, plane below, erect to in- divergentibus vel parallelicis, saepissime flexed above, very often with incomplete, iteratim ramificatis et in cellulas laminae multistratose limbidia; lamina cells inconspicue confluentibus, ultramedio vel irregulary uni- to multistratose, with prope folii extremitatem evanescentibus frequent unistratose areas and patches in exarata, limbis imperfectis, crassis, the upper part, giving the leaf surface an polystratosis praedita. Inflorescentia ut uneven and ragged appearance, smooth sporophyta ignota. or slightly prorulose, thick-walled and porose almost throughout, linear- moderately robust, in lustrous, dense hexagonal to oblong-rhomboidal, straight tufts, typically stiff, fragile and wiry in to somewhat flexuose, attenuate or oblique texture when dry, light to yellowish-green at the ends, 50-90 µm long, 7.5-10.5 µm above, brown to blackish-brown and wide, becoming shorter, rhomboidal, 20- usually incrusted with silt below. Stems 40 µm long at the extreme apex and up to 13 or more cm long, erect-ascending, narrower, 5-7 µm wide, at the margins; sometimes prostrate to suberect, not cells at the insertion uni- to multistratose, radiculose, terete and distinctly catenulate short rectangular, 10-15 µm wide, 15-25 when wet, often attenuate, simple or µm long, with strongly incrassate and sparsely, freely or sometimes fastigiately porose walls, yellow to intensively orange- branched, usually proliferous because of brown on older leaves; alar cells not repeated annual growth from innovations differentiated, short rectangular to formed below the inactive apical cells and subquadrate; costa very difficult for ob- stretching to nearly the same direction as servation and interpretation, imperceptibly that of the preceding stem, light brown merging into multistratose lamina cells, below, green above, in transverse section basically single but more often forked rounded or elliptic, consisting of 2-3 or, in from the large multistratose base occupying places, 4-5 rows of small, rounded cortical 1/3-4/5 the leaf base with 2 main branches, cells with strongly incrassate, light brown divergent or parallel above, ending in the walls surrounding 3-5 rows of large, upper part of the lamina or extending to hyaline, thick-walled medullary cells; the apex, forming distinct crests on the

24 dorsal surface of the leaves and numerous visible in transverse section of the leaves lateral spurs and secondary branches (Fig. 2). Practically, it is impossible to find usually confluent with each other, even two leaves that are similar to one sometimes situated at the margins and another as regards the system of the uni- giving the leaves a limbate appearance, in and multistratose patches and areas in the transverse section composed of small, laminae. As a rule, however, the basal part thick-walled cells of the same shape and of the leaves is 2-5-stratose and can be size as the adjacent lamina cells. No means interpreted as the base of a costa that of asexual reproduction observed. occupies 1/3-4/5 of the leaf insertion. It is Inflorescences and sporophytes unknown. readily observed as an intensively yellow- or orange brown, obscure area Type: Colombia, Dept. Meta: Páramo de imperceptibly merging on one or both Sumapaz, Hoya Sitiales, lagunita 1 km al sides with the very narrow lamina, which E approx. de la Laguna Sitiales, is merely 1-2-stratose and visible as a temporalmente seco, con Myriophyllum more pellucid, yellow to yellowish-green elatinoides y Ranunculus sp., alt. 3650 m, spot. In addition, limbidia frequently occur 22 January 1973, A.M. Cleef 8236 along the leaf margins. These are salient, (Holotype COL; isotypes BM, F, FLAS, swollen, 3-6-stratose leaf borders of H, KRAM, LPB, MEXU, MO, NY, U). varying width and very irregular shape extending from the leaf insertion to mid- leaf or sometimes as far as the leaf apex, where they coalesce with one another as Discussion well as with multistratose streaks radiating Gradsteinia is a very distinct and unique from the costa. genus immediately recognizable from all The costa itself is difficult to interprete other genera of pleurocarpous mosses by because it is not always clearly delimited, its peculiar leaf structure and costa and numerous spurs and secondary condition. The leaves are loosely imbricate, branches radiating from the main branches concave, boat-shaped and cucullate at the or the basal portion of the costa may apex when wet; when dry, however, they considerably obscure its true nature. are curved and inrolled as well as striate Despite these difficulties the costa can be owing to numerous irregular strands and interpreted as basically single but forked crests on their dorsal surface. Young lea- from the base, as seen in the majority of the ves have a short, distinct and mostly leaves. It usually consists of two very recurved apiculus, while the older leaves strong branches, divergent or rarely on the lower part of the stems and branches becoming above, arising from a are without exception fimbriate and incised polystratose base and clearly visible on due to erosion and destruction of their most leaves as distinct, sharply delimited, apices, a feature typical of many aquatic brown crests on their dorsal surface. They and semi-aquatic mosses. However, the cease either freely in the upper part of the most fascinating character of the genus is lamina or extend to the acroscopic margins undoubtedly the state of the lamina cells and fuse with the limbidia, if present, or and the costa. continue their course as marginal or Basically the lamina is unistratose, inframarginal thickenings connivent at the especially distally, but there is an apex. Very often each main branch is extraordinary tendency for multiplication secondarily ramified and spurred, of the number of layers of the lamina cells. sometimes from the base, suggesting a In the absence of any logical rule in the polycostate state of the leaves. However, stratosity pattern, the laminae have a these secondary branches and spurs are as varying and diversified surface as clearly a rule weaker and narrower than the main 25 branches. Because of the frequent reticulate layers of the lamina cells occurs in some connections, the costal system in aquatic genera of the Thamnobryaceae, Gradsteinia is very intricate. Some leaves including Handeliobryum Broth. (Ochyra can in fact may be interpreted as lacking a 1986a), Limbella (C. Muell.) Broth. discrete costa, since it imperceptibly (Ochyra 1987a) and Thamnobryum diffuses into multistratose lamina. Nieuwl. (Ochyra 1990). Although this The peculiar structure of the leaf lamina family is considered to be a hypnobryalean and the costa make Gradsteinia an almost taxon (Buck & Vitt 1986), much evidence unmistakable moss and on that basis alone suggests its closer affinity to isobryalean the genus deserves to be recognized. A neckeraceous mosses rather than to any similar constellation of character states is other group of pleurocarps. All of these unknown in any presently recognized taxa are characterized by having a single genus. The determination of its correct costa and short lamina cells, at least above. systematic position, however, is hampered Because of its elongate lamina cells and by the absence of sporophytes. As forked costa, Gradsteinia could hardly be gametophytic convergence - i.e. the placed in any family of the Leucodonta- occurrence of the same structural character les. A relationship of Gradsteinia to the states in taxa of remote phylogenetic order Hookeriales has been seriously taken relationship - is common in aquatic mosses, into account in my initial considerations. the evaluation of the affinity of Gradsteinia This large order includes many genera remains speculative. and hundreds of species that are mainly Although perichaetia have not been found distributed in the tropics and temperate in Gradsteinia andicola, the presence of areas in the Southern Hemisphere. Its profusely branched stems as well as classification is still debatable and although numerous arrested branch primordia several systems have recently been indicate that this moss is pleurocarpous. proposed (Miller 1971; Crosby 1974; Buck Therefore, considerations on its systematic 1987, 1988; Whittemore & Allen 1989), position should focus on the diplolepidous no general consensus has so far been mosses, especially the orders reached as regard the circumscription of Leucodontales, Hookeriales and families and genera. The main reason of , which are exclusively the incongruity of the various pleurocarpous (Buck & Vitt 1986). classifications of the Hookeriales is the Gradsteinia seems unrelated to any family general lack of correlation between game- or genus of the Leucodontales. This order tophytic and sporophytic characters. As a includes taxa with either thick-walled and result the concept of family in this order isodiametric lamina cells or elongate cells; depends on the importance attributed by when elongate, the leaves are uni- or the autor to characters of either generation ecostate and in addition, have distinct alar Gradsteinia displays some caracters to cells. Polystratosity of the lamina is very support placement in this order, including rarely found in this order, and among bicellular axillary hairs, porose and thick- aquatic genera of this group it is seen only walled lamina cells as well as the lack of in species of Neckeropsis Reichardt sect. pseudoparaphyllia, central strand and Pseudoparaphysanthus (Broth.) Fleisch. differentiated alar cells. The costa condi- (Ochyra & Enroth 1989), Rhabdodontium tion and variously polystratose lamina of Broth. (Norris & Montalvo 1981), Gradsteinia, however, would seem to Muellerobryum Fleisch. or in the very preclude such placement. Apart fron a poorly known Indian moss Pinnatella few ecostate genera such as Stenodesmus limbata Dixon (Dixon 1921), which most (Mitt.) Jaeg. and Phylophyllum C.Muell., probably represents a separate genus. As the Hookeriales include either unicostate well, a tendency for multiplication of the or bicostate genera. In bicostate taxa the 26 costa is perfectly double, i.e. consisting of ron seems rather incongruous in the Hoo- two independent branches arising from keriales; by its leaves it resembles very the leaf base. In some taxa, however, for closely some species of the former genus instance in certain species of Callicosta C. Sciaromium (Mitt.) Mitt. (Ochyra 1987b). Muell. (Crosby 1969), the costae coalesce Therefore, only sporophytic character below and as a result such leaves can be might definitely refute an assumption on interpreted as having single, branched or the relationship of Gradsteinia with forked costae. Such a situation is not hookeriaceous mosses. exceptional in mosses and in several other Partial or perfect polystratosity of the genera with long double costae, for laminae is most often found in the example in Plagiothecium B.S.G., one Hypnales and it is an important diagnostic can observe fusion of the basal part of the character of the Vittiaceae (Ochyra 1987c), costae. Whether the costa in Gradsteinia Hipnobartlettiaceae (Ochyra 1987d) and should be interpreted as a single costa, Donrichardsiaceae (Ochyra 1986b), which which is secondarily forked, or as a double are segregates of the large and very costa wich is secondarily fused at the base heterogeneous family Amblystegiaceae remains a question for discussion. In many and comprize almost exclusively aquatics. leaves of G. andicola the costa consists of It is interesting to note these are unicostate two principals branches that are much taxa, multistratose laminae so far being stronger than the remaining secondary unknown in bi- or ecostate families of the branches and spurs. The two branches are Hypnales including Entodontaceae, divergent or rarely parallel above and Hylocomiaceae, Sematophyllaceae, clearly visible as sharply delimited crests Plagiotheciaceae and Hypnaceae. It is thus on the dorsal surface of the leaves. This obvious the above three families of the situation to some extent resembles that in Hypnales, along with the genus many hookeriaceous taxa with true double Hygrohypnum Lindb. (Amblystegiaceae), costae. which includes many species with spurred As far as I know, multistratose laminae or bifurcate costase, need a careful have not been described for any hookerioid assessment when considering the rela- moss. However, examination of tionship of Gradsteinia. Diploneuron Bartr., a genus endemic to When observing the leaves of G. andicola the West Indies, revealed that Bartram for the first time, I immediately associated (1936) and Allen and Crosby (1986) this moss with Ochyraea tatrensis Vána overlooked partial bistratosity of the (Hypnobartlettiaceae), a remarkable laminae in D. connivens Bartr., the only aquatic pleurocarp endemic to the species of the genus. Diploneuron has a Carpathians (Vána 1986). Like the double costa with two salient branches Colombian Gradsteinia, the European that are parallel after diverging near the Ochyraea has oblong-ovate to oblong- base and continue from about mid-leaf as lanceolate leaves that are concave, entire, limbidia along the acroscopic margins, rounded at the apex and lack a discrete becoming connivent at the apex. The costa. The internal part of the leaves in O. lamina is usually irregularly bistratose, tatrensis is irregularly 2-4-stratose and especially distally, but also in the lower can be interpreted as a costa. It is ill- part of the leaf. Partial bistratosity of the defined, occasionaly interspersed with lamina cells in Diploneuron does not unistratose spots, and imperceptibly suggest the genus should be considered merges with the narrow, unistratose lamina closely related to Gradsteinia but indicates in the fringes of the leaf. The costa of that a tendency for multiplication of the Ochyraea to some extent resembles that layers of the lamina cells does occur in the of Gradsteinia. However, the latter genus Hookeriales. In the sterile state, Diploneu- lacks paraphyllia, while the Ochyraea is 27 characterized by having numerous their misplacement in this genus. They are filamentous paraphyllia on the surfase of either identical to, or supposedly the stem and branches. Because the conspecific with species of Sematophyl- presence and the shape of paraphyllia lum Mitt., Pseudocalliergon (Limpr.) seem to be very indicative for phylogenetic Loeske, Rhynchostegium B.S.G., relationships in mosses, Gradsteinia and Drepanocladus (C. Muell.) Roth, Ochyraea are rather distantly related taxa, Trachyphyllum Gepp, Chryso-hypnum despite the striking similarity of the leaves Hampe and even Bryum Hedw., and bear and costa condition. For the same reason no resemblance to G. andicola, perhaps it is necessary to exclude any relationship except for H. peruviense R. S. Williams. of Gradsteinia to other genera of Hypno- The latter species has broadly ovate, bartlettiaceae, including Hypnobartlettia acuminate leaves with a strong, broad Ochyra of New Zealand, Koponenia costa that is basically single but usually Ochyra of Bolivia, Platylomella Andrews forked or repeatedly divided into 3-5 of eastern North America and branches. Although it is known only in a Cratoneuropsis (Broth.) Fleisch. of sterile state, its gametophyte fits well the Australasia, all having filiform paraphyllia Platyhypnidium-Rhynchostegium (Ochyra 1987d). complex of the Brachytheciaceae. Gradstenia andicola is also entirely Therefore I transferred this species to different from Vittia pachyloma (Mont.) Rhynchostegium as R. peruvianum (R. S. Ochyra, which is the only species of the Williams) Ochyra (in Schultze-Motel & monogeneric family Vittiaceae (Ochyra Menzel 1987). 1987c). This taxon has a single, clearly Until fertile material of Gradsteinia is delimited costa, which is confluent at the available, the best placement for this genus apex with the salient limbidia, while the appears to be in the Donrichardsiaceae. lamina cells are short, rhombic to oblong- Originally this family was established to rhomboidal, mostly unistratose but in some accommodate the monospecific genus populations partially to entirely 2-4-stra- Donrichardsia Crum & Anderson of tose. Such a combination of gametophytic eastern North America. Its principal characters precludes a relationship of Vittia diagnostic characters were a variously and Gradsteinia. multistratose leaf lamina, prorate leaf cells It would be tempting to see a likeness and a very strong and broad costa (Ochyra between G. andicola and some species of 1985). Subsequently Richardsiopsis Hygrohypnum Lindb. of the Ochyra, Sciaromiopsis Broth. and Amblystegiaceae. This genus is extremely Sciaromiella Ochyra have been placed in difficult to define and seems to be a this family (Ochyra 1986b,c). In the present convenient repository for a variety of circumscription the Donrichardsiaceae aquatic pleurocarps with a very variable seems to be unnatural and polyphyletic. costa. As presently defined (Jamieson Unfortunately, the sterile condition of all 1976), Hygrohypnum consist of about 16 genera placed in the family impedes an species that are widely distributed but assessment of their true relationships within scattered in the mountains througout the the Amblystegiaceae-Brachytheciaceae Holartic. No less than 13 species have complex. been described in or transferred to Gametophytically, Gradsteinia is very Hygrohypnum in Central and South unlike Sciaromiella and Sciaromiopsis. America (Wijk et al. 1962; Bartram 1965; Both genera differ from Gradsteinia by Sharp 1978; Crum 1985; Nishimura the presence of very discrete 2-5-stratose 1985). Having examined all relevant type limbidia which are confluent with a single collections of the neotropical costa, as well as ovate-lanceolate and plane Hygrohypnum species, I am confident of leaves. The leaf laminae are irregularly 2- 28 3-stratose in places because of the frequent Crum and Anderson (1979) strongly spurs radiating from the limbidia and, more advocated the relationship of rarely, from the costa. As well, the presence Donrichardsia to Hygrohypnum, but I of pseudoparaphyllia and non-porose feel that suggestion of its alliance with lamina cells seem to preclude a close rela- eurhynchoid genera (Crum 1969) should tionship of these genera with Gradsteinia. be seriously re-assessed. On the other The genus is also very different hand, many species of Hygrohypnum seem gametophytically from Richardsiopsis to have much more in common with some lacustris (Rich. & Herz.) Ochyra of South taxa of the Platyhypnidium- America, which has narrowly lanceolate Rhynchostegium complex of the and gradually long-acuminate leaves and Brachytheciaceae than with other taxa a very strong and long excurrent costa, currently placed in the Amblystegiaceae. which is occasionally slightly spurred Taking into account the aquatic nature of below but otherwise very sharply delimited these taxa resulting in a strong tendency from the lamina cells. for ramification of the costa, Gradsteinia Of all genera currently placed in the would not appear to be anomalous in this Donrichardsiaceae, Gradsteinia seems to group of genera. Therefore its tentative be closest to Donrichardsia. Both genera placement in the Donrichardsiaceae, in are similar in habit, leaf shape as well as the proximity of Donrichardsia, seems to the strong tendency for multistratosity of be acceptable until the interrelationships leaf laminae and branching costae. of some species of Hygrohypnum, Rhyn- However, closer examination reveals chostegium, Eurhynchium, many dissimilarities and differences, which Platyhypnidium and the Donrichardsia- preclude congenericity of these taxa. ceae is critically evaluated. Donrichardsia has (1) broadly foliose pseu- doparaphyllia (none in Gradsteinia); (2) a Acknowledgments small central strand in the stem (none); (3) I am indebted to S. Rob Gradstein for the loan of this interesting thin-walled and non-porose lamina cells material, to Claritza Gradstein-Serna for retyping the manuscript (thick-walled and porose); (4) distinctly and to Bob Magill for language corrections. I am also grateful to prorate cells (smooth or very slightly anonymous reviewers for important suggestions. My wife Halina prorulose); (5) leaves plane (deeply assisted by Miss Katarzyna Noga completed the illustrations and concave and cucullate at the apex); (6) leaf I gratefully ac-knowledge their help. margins sharply serrullate to serrate above (entire or minutely and distantly serrulate). References The costa in Donrichardsia is often laterally spurred but the spurs are generally few Allen, B.H. & Crosby, M. R. (1986). A revision of the and slender. As a consequence, the costa genera Pilotrichidium and Diploneuron (Musci: of Donrichardsia is very distinct and Hookeriaceae). J. Hattori Bot. Lab. 61: 45-64. sharply delimited from the lamina cells Bartram, E.B. (1936). New and noteworthy mosses from contrary to the blurring and unclear costa Jamaica. J. Washington Acad. Sci. 26: 6-l6. in Gradsteinia which gradually merges —— (1965). New and noteworthy mosses from northern into the lamina cells. Likewise, Argentina. Rev. Bryol. Lichénol. 33: 323-327. multistratose lamina strands are clearly Buck, W.R. (1987). Taxonomic and nomenclatural rearran- delimited in Donrichardsia and the large gements in the Hookeriales with notes on the West patches of multistratose cells caracteristic Indian taxa. Brittonia 39: 2l0-224. of Gradsteinia are totally lacking in the —— (1988). Another view of familial delimitation in the Hookeriales. lamina of Donrichardsia. The combination J. Hattori Bot. Lab. 64: 29-36. of these features seems to be sufficient to —— & Vitt, D. H. (1986). Suggestions for a new familial support the generic distinctiveness of classification of pleurocarpous mosses. Taxon 35: 2l- Donrichardsia and Gradsteinia. 60. 29

Crosby, M.R. (1974). Toward a revised classification of the Hattori Bot. Lab. 68: in press. Hookeriaceae (Musci). J. Hattori Bot. Lab. 38: 129- —— & Enroth, J. (1989). Neckeropsis touwii (Musci, Necke- 141. raceae), a new species from Papua New Guinea, —— (1969). A revision of the tropical American moss genus with an evaluation of sect. Pseudoparaphysanthus of Pilotrichum. Bryologist 72: 275-343. Neckeropsis. Ann. Bot. Fennici 26: 127-132. Crum, H. (1969). Nomenclatural notes on North American Schultze-Motel, W. & Menzel, M. (1987). Die Laub- mosses. Bryologist 72: 240-246. moosflora im BRYOTROP-Transekt von Peru. Beih. —— (1985). Two undescribed species of Hygrohypnum from Nova Hedwigia 88: 9-59. Mexico. Bryologist 88: 22-23. Sharp, A. J. (1978). Hygrohypnum steerei sp. nov. from —— & Anderson, L. E. (1979). Donrichardsia, a new genus Mexico. Bryologist 81: 319-321. of Amblystegiaceae (Musci). Fieldiana Botany n. Vána, J. (1986). Ochyraea tatrensis gen. et spec. nov., a ser. 1: 1-8. remarkable pleurocarpous moss from Czechoslo- Dixon, H.N. (1921). On a collection of mosses from the vakia. J. Bryol. l4: 261-267. Kanara District. J. Indian Bot. 2: 174-188. Vitt, D.H. (1984). Classification of Bryophyta. In: Schuster, Jamieson, D.W. (1976). A monograph of the genus R. M. (ed.), New Manual of Bryology, vol. 2, p. 696- Hygrohypnum Lindb. (Musci). 425 pp. Ph. D. thesis, 759. Nichinan. Univ. of British Columbia, Vancouver. Whittemore, A. & Allen, B. H. (1989). The systematic Miller, H.A. (1971). An overview of the Hookeriales. position of Adelothecium Mitt. and the familial Phytologia 21: 243-252. classification of the Hookeriales (Musci). Bryologist Nishimura, N. (1985). A revision of the genus Ctenidium 92: 261-272. (Musci). J. Hattori Bot. Lab. 58: 1-82. Wijk, R. van der, Margadant, W. D. & Florschütz, P. Norris, D. H. & Montalvo, A. M. (1981). The rediscovery A. (1962). Index Muscorum, Vol. 2 (D-Hypno). of Rhabdodontium buftonii (Broth. & Geh. in Broth.) Regn. Veget. 26: 1-535. Utrecht. Broth. Bryologist 84: 85-88. Ochyra, R. (1985). Hypnobartlettia fontana gen. et sp. nov. (Musci: Hypnobarlettiaceae fam. nov.), a unique moss from New Zealand. Lindbergia 11: 2-8. —— (1986a). A taxonomic study of the genus Handeliobryum Broth (Musci, Thamnobryaceae). J. Hattori Bot. Lab. 61: 65-74. —— (1986b). Sciaromiadelphus A. Abr. & I. Abr. - the relationship between extant and fossil moss specimens. J. Hattori Bot. Lab. 61: 309-332. —— (1986c). On the taxonomic position of Sciaromium lacustre Herz. & Rich. J. Bryol. 14: 109-115. —— (1987a) On the and family placement of the moss genus Limbella (C. Muell.) Broth. J. Bryol. 14: 465- 485. —— (1987b) A revision of the moss genus Sciaromium (Mitt.) Mitt. - corollarium. Bull. Polish Acad. Sci. Biol. Sc. 35: 33-45. —— (1987c). A revision of the moss genus Sciaromium (Mitt.) Mitt. II. The section Limbidium Dusén, with a description of Vittia gen. nov. (Vittiaceae fam. nov.). J. Hattori Bot. Lab. 62: 387-415. —— (1987d). A revision of the moss genus Sciaromium (Mitt.) Mitt. III. The section Platyloma Broth. J. Hattori Bot. Lab. 63: l07-132. —— (1990). On the relationship of Thamnobryum negrosense (Bartr.) Iwats. & Tan (Musci: Thamnobryaceae). J.