Marsupella tubulosa Steph., Bull. Herb. Boiss. 5: 99, 1897

Bakalin, Vadim, Choi, Seung Se & Park, Seung Jin, 2021, Revision of Gymnomitriaceae (Marchantiophyta) in the Korean Peninsula, PhytoKeys 176, pp. 77-110 : 77

Marsupella tubulosa Steph., Bull. Herb. Boiss. 5: 99, 1897 Figure 5

Marsupella emarginata var. tubulosa (Steph.) N.Kitag., Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Biol. 27: 77, 1960.

Type.

Japan, Unzen , 5 Mar 1895, Faurie 15385 (lectotype (designated here) G [15042/00061032!])

Description.

Plants merely rigid, forming loose patches, deep green-brown, purple-brown, purple-green, or rarely greenish (actually plants extracted from the patch yellowish brownish in general, but with purple-rusty coloration in apices and upper parts of insolated leaves that gives expression of purple-brown color of patch), or yellowish greenish, pale brownish with purple tint in apical parts, rarer brownish greenish without purple or rusty pigmentation; 0.6-1.2 mm wide (the largest lax plants up to 1.3-1.5 mm) and 5.0-15.0 mm long. Rhizoids nearly absent or few, colorless. Stem brownish, not branched or branched as ventral leafless stolons (rarely becoming to normal branch) or more commonly as subfloral ventral or lateral innovations; transversely elliptic in cross section, 160.0-180.0 μm high and 200.0-250.0 μm wide (depauperate shoots omitted), differentiated into strata; hyaloderm with external wall thin, radial walls thin to unequally thickened (becoming thicker inward), inner wall thick, 15.0-20.0 μm along margin; scleroderm in (1-)2 rows of cells, cells thick-walled, but not so strongly as in M. apertifolia ; inner cells thin to slightly thickened, trigones moderate in size, concave, 12.0-18.0 μm in diameter. Leaves contiguous to distant, sometimes ‘enclosed’ one to another, concave to almost flattened in upper half, transversely inserted, evidently or very loosely sheathing stem in the base, obliquely (rarely erect) spreading, subtransversely to obliquely oriented, suborbicular to widely ovate, margin flat to loosely recurved near base (especially in the leaves sheathing the stem), divided into two unequal (rarely subequal) lobes by widely V- to γ -shaped sinus descending to 1/7-1/5(1/4) of leaf length, lobe apex couple-shaped, obtuse to acute. Cells in the midleaf shortly oblong, 10.0-20.0(-25.0) × 8.0-16.0(-18.0) μm, walls thin to slightly thickened, trigones large to (rarer) moderate in size, convex to bulging, cuticle smooth; cells along leaf margin 6.0-12.0 μm, walls thin to slightly (to strongly and unequally) thickened in tangential walls, trigones moderate to large, concave to slightly convex, tangentially sometimes confluent; cells in lobe middle oblong, 15.0-22.0 × 8.0-13.0 μm, walls thin to slightly thickened, trigones moderate to large, convex to triangle. Oil-bodies in the midleaf cells 2(-3) per cell, biconcentric (at least (30-)70%), finely to normally papillose, spherical, ca. 5.0 μm in diameter to oblong, 7.5-12.5 × 5.0-7.5 μm. Dioicous. Androecia intercalary, with 1-3 or 3-5 pairs of bracts, different generations divided by 4-8 pairs of sterile leaves, spicate, 3-5-androus, stalk biseriate, 75.0-100.0 μm long, body ellipsoidal, ca. 140.0 × 115.0 μm; bracts spoon-shaped, nearly subquadrate when flattened, with deflexed to erect-spreading lobes. Perianth onion-shaped, hidden within bracts, ca. 300.0 × 500.0 μm; perigynium 500.0-700.0 μm long, with (1-)2 pairs of bracts; bracts sheathing perianth near base and erect spreading in upper 1/3-1/2. Capsule elliptic, outer layer cells rectangular, 27.0-50.0 × 20.0-25 μm, with 2(-4) nodular thickenings in vertical walls and 1(-3) thickenings in horizontal wall; inner cells elongate and flexuous, 45.0-63.0 × 7.0-10.0 μm, with 7-10 sometimes bifurcate semicircular bands. Elaters (2-)3-spiral, 150.0-180.0 × 7.0-8.0 μm. Spores brown, papillose, spherical, 10.0-12.0 μm in diameter.

Ecology.

Acidophilic meso- to hygrophyte. The species occupies sandy soils and mineral substrates, over wet to moist, and sometimes mesic cliffs, being most common along streams near running water. In drier habitats, it is commonly associated with Odontoschisma pseudogrossiverrucosum , Cheilolejeunea obtusifolia , and rarely with Microlejeunea punctiformis , Cephaloziella spp., Gymnomitrion faurianum . In wetter habitats M. tubulosa sometimes grows intermixed with Solenostoma minutissimum , Lophocolea horikowana , Marsupella pseudofunckii , and M. koreana .

Distribution.

The distribution of the species is confined to insular and peninsular areas in Amphi-Pacific Boreal and Temperate Eastern Asia, while the records provided by Bakalin, (2010), Cherdantseva and Gambaryan (1986), Gambaryan (1992, 2001), Konstantinova et al. (2002) for the Russian Far East continental mainland are likely incorrect. This species is strikingly characterized by biconcentric oil bodies - an uncommon feature in Marsupella that has never been reported from continental Asia or from North America. This suggests the reports of the species in Schuster (1974) for North America and Schljakov (1981) for Russian Asia are incorrect. We suggest the ‘true’ M. tubulosa (the type is from Honshu) occurs only in Japan, Kurils, Kamchatka and the Korean Peninsula, as well as probably in China (Anhui, Taiwan), from where, unfortunately, oil bodies were not studied. All specimens collected in the continental mainland of the Russian Far East and checked alive had non-biconcentric oil bodies, whereas specimens from peninsular and insular parts of the Far East possess biconcentric oil bodies. The species was recorded for nearly all provinces of the Korean Peninsula (Jeju-do, Gyeongsangnam-do, Gyeongsangbuk-do, Chungcheongnam-do, Chungcheongbuk-do, Gyeonggi-do, Gangwon-do, Pyeonganbuk-do, Hamgyeongnam-do, Hamgyeongbuk-do: Yamada and Choe 1997; Kim and Hwang 1991) and was confirmed for most of the provinces in the southern part of the peninsula.

Specimens examined.

Chungcheongnam-do: Mt. Daedun , 36°08'02.9"N, 127°18'29.1"E, 343 m, 31 Mar 2009, S.S. Choi 3399 (JNU), Mt. Gyeryong , 36°21'06.9"N, 127°12'50.9"E, 290 m, 8 Jul 2009, S.S. Choi 4098 (JNU); Gangwon-do: Mt. Seorak , 38°06'02.8"N, 128°23'43.1"E, 840 m, 28 Aug 2009, S.S. Choi 4258 (JNU), Mt. Seorak , 38°06'39.7"N, 128°24'56.2"E, 1347 m, 21 Sep 2009, S.S. Choi 5095 (JNU), Mt. Seorak , 38°07'21.0"N, 128°27'27.7"E, 1649 m, 21 Sep 2009, S.S. Choi 5175 (JNU); Gyeongsangbuk-do: Is. Ulleung, Seonginbong, 37°29'39.5"N, 130°52'35.4"E, 845 m, 20 Oct 2010, S.S. Choi 8712 (JNU), 37°29'53.9"N, 130°52'01.0"E, 977 m, 20 Oct 2010, S.S. Choi 8744 (JNU); Gyeongsangnam-do: Mt, Gaya, 35°48'53.9"N, 128°07'21.9"E, 1116 m, 8 Sep 2009, S.S. Choi 4361 (JNU), 35°49'14.8"N, 128°07'27.5"E, 1313 m, 8 Sep 2009, S.S. Choi 4376 (JNU), Mt. Jiri , 35°18'51.9"N, 127°44'22.1"E, 848 m, 13 Jun 2009, S.S. Choi 3641 (JNU), 35°19'50.7"N, 127°44'08.1"E, 1540 m, 14 Jun 2009, S.S. Choi 3732 (JNU); Jeju-do: Bolre Oreum, 33°21'20.8"N, 126°28'10.2"E, 1145 m, 5 Sep 2012, S.S. Choi 120721 (JNU), Erimok valley , 33°21'59.6"N, 126°30'40.25"E, 1613 m, 6 Sep 2012, S.S. Choi 20795 (JNU), Mt. Halla , 33°21'43.1"N, 126°32'21.9"E, 1835 m, 8 Aug 2010, S.S. Choi 7745 (JNU), Musu stream, 33°25'08.0"N, 126°26'56.4"E, 495 m, 28 Oct 2010, S.S. Choi 8827 (JNU), Suak valley , 33°20'14.0"N, 126°36'37.8"E, 523 m, 29 Oct 2010, S.S. Choi 8879 (JNU); Jeollabuk-do: Mt. Deogyu , 30 Jun 2008, S.S. Choi 887 (JNU), Mt. Jiri , 35°19'06.1"N, 127°31'47.5"E, 781 m, 20 Jun 2009, S.S. Choi 3993 (JNU); Jeollanam-do: Mt. Dureun, 5 Feb 2009, S.S. Choi 3061 (JNU), Mt. Jogyeo, 34°59'51.6"N, 127°16'51.7"E, 288 m, 7 Dec 2010, S.S. Choi 9094 (JNU) .

Comments.

This species may be confused with at least three other species: Marsupella koreana , M. apertifolia , and M. emarginata , though the last is not known from the Korean Peninsula. The distinctions between the former two are mentioned under those species. The species differs from M. emarginata (with which M. tubulosa was probably confused in parts outside of oceanic/suboceanic Eastern Asia) in biconcentric oil bodies (versus oil bodies without the central eye) and commonly obliquely oriented leaves that are not closely sheathing the stem in the base (versus transversely oriented leaves with leaves closely sheathing the stem near base). The latter feature is surely quantitative and variation in leaf orientation occurs within the two species.

Figures

Figure 5. Marsupella tubulosa Steph. A gynoecium longitudinal section B stem cross section (fragment) C plant habit, dorsal view D leaf margin cells E, F leaves G elater, H midleaf cells. Scale bars: 1 mm (A, C, E, F); b 100 µm (B, D, G, H). All from Choi 3732 (JNU).