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Studieson Leacodon (Leucodontaceae

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Studieson Leacodon (Leucodontaceae The JapaneseSocietyJapanese Society for Plant Systematics Plrytotax. June, 19ee Acta Geobot. 73 Studies on Leacodon (Leucodontaceae, Musci) and related genera in East Asia III*. Notes on the systematic position oi PTerogonium Sw. Hiroyukl AmyAMA*" : -" -" llkilagk2 M7 i> 7gi B f b're (l i9 iF b-isF,ptff)rks.k as 4(DiEEwakOiifiscM, Pterogonium tc, ･eafitw\ftSl twaxSUICOUi-Owtst Since BRucH et al. (1851) chose the leetotype of the genus, the generic composition of Pterqgenium has been refined. Pteragonium is at accepted as monotypic comprising , present only one species, P. gracile (HEDw.) SM. (WiJK et al. 1967). There are several opinions on the systematic position of Pterqgonium. BRuaH et al. (1851) and ScHiMpER (1855) considered it as a member of the Pylaisaeaceae because of its erect capsule and pleurocarpous habit. But, nowaday their family concept appears to be a mere assemblage based on a superficial resemblance and so, quite unnatural. established the Pterogoniaceae to accomodate Pterogonium and g ScHiMpER (1860) Ptetig)nandntm. He listed several characteristics for this family: (i) the presenCe of ascending shoots; (ii) the simple er forked costae; (iii) the smooth or dorsally paplllose Ieaves; (iv) the presence of its symmetric and erect capsules, cucullate calyptrae and he transferred numerous paraphyllia; (v) the vestigial peristomes. But later (1876) Pteragonium in the Leucodontaceae because of rhomboidal to linear laminal cells. LiMpRiaHT (1895) included Pteregonium in the Leskeaceae s. Iat. on the basis ofthe papillae on laminal cells and the forked costae; he did not distinguish papillae on cell lumens frorn fbllowed SaHiMpER prorations situated at the upper corner of cells. BRoTHERus (1905) (1876) and placed Pteregonium in the Leucodontaceae. should be included Until now, almost all bryologists have considered that PterQgonium 1954, in the Leucodontaceae (BRoTHERus 1925, FLEiscHER 1906, DixoN 1924, NyHoLM LAwroN 1971, ViTT 1984, and many others). However, the position and relationships of Pterogonium within the family has been controversial. FLEiscHER (l906) proposed a general system ofmoss classificatign accepted by most "lacked" later bryologists. He pointed out that Pteragonium costa and had a unique * continued from Bryologist 90: 4e9--416. (1988). ** DepaTtment of Botany, Faculty of Science, Kyoto University, KyoTo 606, Japan NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics 74 Phytotax. Acta Geobot. Vol. XXXIX, Nos: 1-3 similar peristome to that of Lettcodbn, and assigned Pteragonium in the Leucodontaceae. But,because of itsdistinctivenesswithin the family, ' he placed Pterrgonium in a new tribe efits own, Pterogoneae. BRoTHERus (1925) foIIowed FLEiscHER but he raised the tribe ' to the rank of subfamily in the Leucodontaceae. MANuEL (1974) divided the Leucodontaceae into four subfamilies on the basis of three charabters: gametophytic presenc.e or absence of paraphyllia, pseudoparaphyllia, and a central strand in the stem. He was the first to point out the close relationship between Pteragonium, Antitriehia, and Bestia. He placed these genera in the Pterogoneae in the Leucodontoideae. According to him, these three genera share tlte fbllowing features: the exostome teeth horizontaily (i) striated below and smooth or papillose above; the well (ii) developed end6stome. ' BucK adopted MANuEL'sinfrafamilial (1980a) classification. He, however, proposed to transfer the above three genera together with another genus, Atsia, from the Leucodont- aceae to a family distinct Pterogoniaceae. He pointed out that these four genera were closely related cach othcr in sharing (i) the short, thick-walled laminal cells, (ii) the costae frequently forked at base into 2-3 main branches, (iii) the shelfilike habit, (iv) the short emserted capsules, and (v) the peristome papMose or with a hint ofstriation on the base of the exostome teeth. Any treatments mentioned above, however, do not seem to be based on adequate morphological observations ofboth gametophytes and sporophytes. In order to clarify the systematic positien of Pteragonium, re-examination ef them are needed not only for Pteragonium itselfbut also fbr the genera hitherto considered to be closely related to it. ' ' ' h. Observations Gametophytes of Ptengonitim 1. Habit ' Plants consist of three parts, i.e., stoloniform shoots, ascending shoots and flagella. 'and Stoleniform shoots prostrate on substratum (tree trunk or rock) branch irregularly. Ascending shbots consist of lower parts forming stipes and of upper parts with many branches densely and pinnately arranged. Each branch is at a right angle to the main a#is of the ascending shoet. Tips ef branches and main axis are often fiagelliform. They never break away as in the case ef brood branchlets often observed in the Leucodontaceae. Ascending shoots showjulaceeus habit. Branches curve on one side, and so plants show characteristic appearance in dry conditions. ' ' 2. Leaves Distinct differentiationsoccur in shape and size among stipe, stem, and branch leaves. Stipe leaves have cordate bases which are abruptly narrowed into slender upper 'Laminal parts. They are appressed to the stipe even in wet conditions. cells of stipe leaves are slightly longer than those of stem and branch !eaves. Serration of upper NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics June,1988 Acta Phytotax. Geobot. 75 18 Fig. I. 1-16. Pterogonium gracile. 17. Antitrichia curtipendula. 18. Antitrichia catijbrnica. 1, stem leaf <x25); 2, branch leaf(× 25)l 3, stipe leaf(x25); 4, tip of stem leaf (× 270); 5, upper 1aminal cells with distinct proration C× 270) ; 6, median Iam{nal cells C × 270) ; 7, upper margin of stem leaf ( x270) ; 8, lewer margin of stem leaf C× 27e) ; 9, scale-like leaf of branch primordium (x270); 10 and 11, axillary hairs (× 270); 12, pTant in wet condition (× 3); 13, capsule (× 7); 14, spores (x270); 15, leaf( perichaetial × 25); 16, transverse section of stem (× 270); 17 and 18, stem leayes (× 25), l-10 and 12-16, S2:hofieid 29014 (NICH). 11, Revik s.n. (NICH). 17, Sbhqi7eldet al. 43896 (HIRO). 18, Sthojield 284s3 (HIRO). NII-Electronic Library Service The JapaneseSocietyJapanese Society forforPlant Plant Systematics 76 Acta Phytotax. Geobot Vol. XXXIX, Nos. 1-3 ' margins is low. Stem leaves are ovate-lanceolate with distinct serration (Fig. I 4, 7); each tooth is composed of l-3 cells. Branch leaves are similar to stem leaves but usually much smaller. In spite of these diflerences of stipe, stern, and branch leaves, the fo11owing features are shared among them; (i) the proration at the upper corner of several cells, (ii) the presence of serration, (iii) the distinct alar region composed of quadrate cells, (iv) the forked costa with several short protrusions in the basal part. The degree of proration varies among specimens and in a few case no proration is observed. 3. stems ' ' more rounded transverse sections and the Stems of ascending shoots are or less in constituent cells are diff;:rentiated into three layers ; epidermis, cortical layer and a central strand (Fig. I 16). Paraphyllia and pseudoparaphyllia are absent; branch primordia are covered by smal}, scale-like leaves which are derived from the branch primordium high) itself (Fig. I 9). Axillary hairs consist ofbrownish, quadrate basa! cells (1-2 cells and colorless, eblong upper cells (1-3 cells high) ; the hight varies among specimens. ' ' 4. Perichaetial leaves ' stem leaves. Perichaetial leaves lengthen after fertilization and are much longer than There are low serrations oniy at the upper part. Costae are fbrked at base arid lack protrusions in the basal part. Laminal cells are much elongated-rectangular without proration. Hairs arising from vaginula are few to numerous and usually uniseriate (partly biseriate). 5. 0thers ' Calyptrae are cucullate and provided with few hairs. Sexuality is dioecious. Fe- male and male plants are nearly ofthe same size. Gametangia are restricted to the main axis of ascending shoot. Sporophytes of Pteragenium The peristorne consists ofexostome and endosteme. The properistome (= prostome) is absent. Each tooth ofexostome is lanceolate, The outer surfaces ofexostome teeth have horizontal er oblique striae at base and dense papillae above (Fig. II 3-zl). These striae are, however, diMcult to observe with a light microscope. The endostome consists of a high basal membrane and short segments (Fig. II 2), which are smooth on both surfaces, lacking any ornamentation. The segments are arranged alternately against exostome teeth. Cillia are lacking. In moist conditions, exostome teeth curve inward. In dry conditions, they spread outward at first and then only their upper parts become incurved IV upper). The ' (Fig. endostome is immobile in any conditions. Capsules are long exserted from perichaetial leaves, cylindric and slight!y narrowed at mouth, smooth (slightly plicate when dry). Stomata are present at apophyses. NII-Electronic Library Service The JapaneseSocietyJapanese Society for PlantPlantSystematics Systematics June, 1988 Acta Phytotax. Geobot. 77 Exothecial cells are rectangular and thick-walled. Speres are spherical, thin- walled, rninutely papshortillose,13-16 pam in diameter, isosporic. Discussion Comparison with Leucodon It seems evident from the present study that the observations by FLEisGHER (1906) on Pteragonium and Leucodbn are incorrect. Indeed, concerning the suggested resemblances in costa and peristome morphology, Pteragenium has
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