Marsupella apertifolia Steph., Bull. Herb. Boiss. ser. 2, 1: 162 (Spec. Hep. 2: 23), 1901

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

Marsupella apertifolia Steph., Bull. Herb. Boiss. ser. 2, 1: 162 (Spec. Hep. 2: 23), 1901 Figure 2

Marsupella emarginata subsp. tubulosa var. apertifolia (Steph.) N.Kitag., J. Hattori Bot. Lab. 26: 89, 1963.

Marsupella tubulosa var. apertifolia (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 78, 1944.

Type.

Japan, Miyokosan, U. Faurie 75 (lectotype (designated here): G [009469!]) .

Description.

Plants in rather loose patches, rigid to more soft, erect or nearly so, mostly deep green to brownish green in color, but with many other variants intergrading to yellowish-brownish, yellowish, and pale greenish (later only in shaded wet places) or to deep brown and rusty pigmentation in insolated moist habitats; mostly 1.2-1.6 mm wide and 7.0-15.0 mm long, with small forms starting from 0.75-1.0 mm wide and robust varying to 1.5-2.2 mm wide. Branching lateral (rare) or ventral as subfloral innovations (more common), stem transversely elliptic in the cross section, 200.0-300.0 μm high (extreme variants not included) and 250.0-350.0 μm wide; outer cells (hyaloderm) nearly thin, with small trigones, 15.0-25.0 μm along margin, scleroderm in 3-4 layers, with cells slightly smaller, very thick-walled, with lumen just 8.0-11.0 μm, inner cells (10.0-)12.0-20.0 μm, thin- to slightly thickened, with moderate in size, triangle to concave trigones. Rhizoids sparse to virtually absent, mostly colorless to brownish, in unclear obliquely to erect spreading fascicles, rarely (and very few in number) separated and deep purple. Leaves mostly contiguous and loosely enclosed one to another, to subimbricate or nearly distant in lax modifications, concave to canaliculate-concave, transversely inserted, evidently sheathing the stem in the base and obliquely to erect spreading above, transversely, subtransversely or (more rarely) obliquely oriented, with margin commonly narrowly recurved, at least in lower half of the leaf; transversely elliptic to orbicular and widely ovate in shape, mostly 500.0-750.0 μm long and 550.0-1050.0 μm wide, reaching in lax forms 1500.0-2250.0 × 1550.0-2750.0 μm, divided by sinus descending to 1/7-1/5(1/4) of leaf length into two nearly equal to subequal lobes; sinus varying from narrowly to widely γ -shaped; lobes rounded to (rarely) obtuse in apex. Cells in the midleaf subisodiametric to shortly oblong, (12.0-)20.0-25.0 × (12.0-)13.0-25.0 μm, thin-walled to slightly thickened, trigones mostly large, rarer moderate in size, convex to bulging, cuticle smooth; cells along margin in upper part of leaf 5.0-8.0(-10.0) μm, mostly unequally thickened due to trigones confluence, trigones large, convex to concave, cuticle smooth; cells in the lobe middle similar to that in the midleaf or slightly smaller, 12.0-20.0 × 11.0-15.0 μm, thin-walled, trigones large, convex to bulging, cuticle smooth. Dioicous. Androecia intercalary, spicate, with 3-4 pairs of bracts, (1-)2-3-androus, antheridium body obovate, 130.0-150.0 μm wide, stalk biseriate, 100-150 μm long; bracts strongly inflate in lower half and obliquely to erect (especially lobes) spreading above, trapezoidal-subtransversely elliptic. Perianth hidden within bracts or very shortly exerted, conical to onion-shaped, 400.0-750.0 × 750.0 μm; perigynium 750.0-1000.0 μm long, with two pairs of bracts; bracts sheathing perigynium in lower part and obliquely spreading above (lobes of lower pair commonly deflexed).

Ecology.

Acidophilic hygro- to hydrophyte, occupying various habitats, from very wet (and even submerged) shaded cliffs near running water to moist mineral substrata in full sun. In moist and sunny habitats, robust phases are formed (then commonly acquiring deep rusty-brown pigmentation), where it is associated with Anastrophyllum assimile , Trilophozia quinquedentata , and Diplophyllum taxifolium . As an extreme variant, the species may be intermixed with Gymnomitrion faurianum . In wet and shady habitats, its common association is Cephalozia otaruensis *.

Distribution.

Montane temperate Kurils-Japanese-Korean endemic species is known in northern and middle Japan (until Shikoku), South Korea and South Kurils (Iturup Island), likely more widely distributed, at least to Kamchatka Peninsula in in the north. In Korea, Jeju-do, Chungcheongbuk-do, Chungchengnam-do, Gyeongsangnam-do, Gangwon-do, Jeollabuk-do and Jeollanam-do ( Choi et al. 2017).

Specimens examined.

Chungcheongnam-do: Mt. Daedun , 36°08'02.9"N, 127°18'29.1"E, 343 m, 31 Mar 2009, S.S. Choi 3405 (JNU); Gangwon-do: Mt. Seorak , 38°07'21.0"N, 128°27'27.7"E, 1649 m, 21 Sep 2009, S.S. Choi 5174 (JNU), Mt. Seorak , 38°07'42.2"N, 128°26'21.6"E, 1011 m, 14 Oct 2010, S.S. Choi 8607 (JNU), Mt. Seorak , 38°07'52.7"N, 128°26'11.2"E, 937 m, 14 Oct 2010, S.S. Choi 8632 (JNU); Gyeongsangnam-do: Mt. Jiri , 35°19'20.6"N, 127°44'59.4"E, 1134 m, 14 Jun 2009, S.S. Choi 3745 (JNU), Mt. Jiri , 35°20'01.7"N, 127°43'55.1"E, 1713 m, 3 Oct 2011, S.S. Choi 111079 (JNU), Mt. Namdeogyu , 30 Oct 2008, S.S. Choi 1119 (JNU); Jeju-do: Erimok valley , 33°21'59.6"N, 126°30'40.3"E, 1591 m, 6 Sep 2012, S.S. Choi 120765 (JNU), Erimok valley , 33°21'59.6"N, 126°30'40.27"E, 1615 m, 6 Sep 2012, S.S. Choi 120797 (JNU), Mt. Halla , 33°22'02.2"N, 126°33'05.9"E, 1563 m, 8 Aug 2010, S.S. Choi 7737 (JNU), Mt. Halla , 33°21'42.1"N, 126°32'02.8"E, 1861 m, 21 Sep 2012, S.S. Choi 120904 (JNU), Hyodon stream, 33°18'21.4"N, 126°33'38.5"E, 469 m, 7 Aug 2010, S.S. Choi 7638 (JNU), Witse Oreum, 33°21'33.4", 126°30'54.2"E, 1668 m, 7 Sep 2012, S.S. Choi 120847 (JNU); Jeollabuk-do: Mt. Deogyu , 22 Nov 2008, S.S. Choi site 2-35 (JNU), Mt. Jiri , 35°19'25.0"N, 127°41'36.8"E, 1300 m, 7 Oct 2009, S.S. Choi 6090 (JNU), Mt. Jiri , 35°19'50.1"N, 127°41'33.5"E, 1100 m, 21 May 2010, S.S. Choi 7383 (JNU); Jeollanam-do: Mt. Dureun, 5 Feb 2009, S.S. Choi 3064 (JNU), Mt. Jiri, 35°19'15.3"N, 127°31'50.0"E, 755 m, 19 Sep 2009, S.S. Choi 5043 (JNU) .

Comments.

This species was regarded as the variety within M. emarginata subsp. tubulosa by Kitagawa (1963); however, we agree with Stephani (1901) and treat it as a separate species because of the differences in DNA sequences between two taxa ( Bakalin et al. 2019). Marsupella apertifolia differs from M. tubulosa in mostly rounded lobe apices (versus mostly acute), more or less equal lobes (versus distinctly unequal), non-biconcentric oil bodies (versus biconcentric), and constant absence of red or purple pigmentation.

Figures

Figure 2. Marsupella apertifolia Steph. A male plant B female plants C stem cross section (fragment) D plant habit E-H leaves I, K cells along leaf margin J midleaf cells L gynoecium longitudinal section. Scale bars: a 1 mm (A, B, D-H, L); b 100 µm (C, I-K). All from Choi 7383 (JNU). Drawing by S. J. Park.