Fabronia ciliaris (Brid.) Brid., Bryol. Univ.

Ignatova, Elena A., Kuznetsova, Oxana I., Milyutina, Irina A., Fedosov, Vladimir E. & Ignatov, Michael S., 2017, The genus Fabronia (Fabroniaceae, Bryophyta) in Russia, Arctoa 26 (1), pp. 11-34 : 26-32

Fabronia ciliaris (Brid.) Brid., Bryol. Univ. 2: 171. 1827.

— Hypnum ciliare Brid., Muscol. Recent. Suppl. 2: 155. 1812.

Holotype. “Schleicher Botaunte Fabronia octoblepharis , Fabronia ciliaris Bryol. Univ. Hypnum [sericeulum?] Helvetia, 1807” (B 31 0677 01), http:// ww2.bgbm.org/Herbarium/specimen.cfm?Barcode= B31067701 ( Röpert, 2000).

Putative isotype in LE: “ Pterigynandrum obctoblepharis Schl. , am Nov. genus?, ex Helvet” [another later hardwriting at label bottom “ Fabronia ”].

Illustrations: Figs. 5 : 1–6, 6: 1–5, 11: 1–7; Ignatov & Ignatova (2004): Fig 429 on page 647.

Plants small, silky-green or yellow-green, glossy, in dense patches. Stems creeping, ca. 5 mm long, irregularly branching; branches 2–3 mm long, erect, densely foliate. Stem and branch leaves similar, loosely appressed when dry, straight, often with slightly recurved apices, spreading when moist, ovate, abruptly long acuminate, 0.5–0.9 u 0.25–0.3 mm; leaf margins denticulate, marginal teeth unicellular, small (ca. 0.1 mm long), or, rarely, consisting of 2–3 cells; costa slender, smooth, ending at midleaf; lamina smooth, upper and median laminal cells rhomboidal, 20–35(–40) u 8–11 µm, with length/width ratio 2–4:1 and cell length, µm/ leaf length, mm ratio 27.0– 53.3; apical cell ca. 110 µm long; alar cells quadrate to short-rectangular, forming weakly delimited rectangular goup 3–4 cells wide and 6–9 cells along leaf margin. Autoicous, sporophytes rare. Perigonia bud-like, sometimes numerous. Perichaetial leaves 0.4–0.5 mm long and 0.3–0.4 mm wide, with oblong base and gradually attenuate narrow acumen; costa ending below apex; margins with longer, stronger recurved teeth in upper part. Sporophytes single in perichaetium. Seta 3–4 mm long, straight, flexuose when dry, yellow. Capsule ovate, with short neck, 0.35–0.45 mm long and 0.3–0.4 mm wide, often flared at mouth with age, smooth, pale brown, concolorous at rim or slightly darker; annulus absent. Peristome single, consisting of 16 exostome teeth fused in pair, light-brown or brown, obtuse, ca. 125 µm long; outer surface papillose-striolate, inner surface with less prominent ornamentation, in distal parts of teeth covered with low vertical ridges. Spores (10–)15–17 µm, papillose. Operculum low conic, mammillate or with short straight or, rarely, moderately long oblique beak.

Distribution and ecology. Fabronia ciliaris s. str. is considered as a widespread species throughout the world. It is known from mountain areas of Central and Southern Europe, in Asia it was reported from many provinces of China, from Korea and Japan; it is quite widespread in America, especially in the eastern part, uncommon in Mexico and central South America and becoming more frequent to the south, in mountain areas ( Buck, 1983); it was also reported from Australia, New Zealand and Hawai. In Russia, F. ciliaris is most widespread and frequent species of the genus. It is known from Eastern and Central Caucasus, Middle and South Urals, southern Siberia and south of Russian Far East; its northernmost records were made in southern and central Yakutia and the extreme point is in Verkhoyansk District, at 67°35’N ( Isakova, 2010). Most numerous collections were made in Altai Mt. and Transbaikalia. This species grows in mountain areas in a wide range of altitudes, from 150 to 1550 m a.s.l., but mainly in forest zone. It inhabits dry and moderately wet cliffs, rock outcrops and boulders, both acidic and calcareous, in open and shady places, and occasionally grows on soil in rocky places; it was also collected on tree trunks: on poplar in Khabarovk Territory, on elm in Primorsky Territory and on pear trees in Dagestan. In Yakutia, it occurs on rock outcrops on xeric and warm steppe slopes, where other southern xerophilous species where found.

Selected specimens examined: RUSSIA: CAUCASUS: Dagestan Republic: Gunib District: Gunib , 1550 m alt., 22. V.2009, Ignatov & Ignatova 09-672 & 09–766 ( MW) ; North Ossetia /Alania: Suargom Gorge, 800 m alt., 1.XII.2014, Ukrainskaya 15945 ( LE) ; Kabardino-Balkaria: Chereksky Disrict, Verkhnaya Balkaria Settl., 1100–1150 m alt., 28.VIII.2005, Ignatov et al. 05-1825 & 05-1880 ( MHA) ; Elbrussky District, Bylym, Baksan River valley , 1000 m alt., 30.VII.2004, Ignatov et al. s.n. ( MHA) ; Karachaevo-Cherkessia: Teberda Nature Reserve, Dzhemagat Creek valley , 1550 m alt., 22.VII.1994, Onipchenko 115/94 ( MW) ; Aksaut Gorge, 5. VI.1989, Ukrainskaya 14444 ( LE) ; Daut River valley, 4.VIII.1993, Ukrainskaya 14443 ( LE). URALS: Bashkortostan: Bashkirsky Nature Reserve , Kaga River mouth, 2.IX.1945, Selivanova-Gorodkova s.n. ( MHA) ; Chelyabinsk Province: Satka District, Ai River, 300 m alt., 6.VIII.2011, Ibatullin 36 ( MHA) ; Perm Territory: Kishertsky District, Kamen Vostryj , 2.VII.1999, Bezgodov 268 ( MW); ASIAN RUSSIA: Altai Republic: Ulagan District: Chulyshman River valley, 1950 m alt., 2.VIII.1993, Ignatov 36/95 ( MHA) ; Chulcha River valley, 1000 m alt., 11.VII.1991, Ignatov 9/12 ( MHA) ; northern shore of Teletzkoe Lake, Yailyu, 460 m alt., 1. VI.1989, Ignatov 0/1054 ( MHA) ; west of Yailyu, Izvestkovaya Mt., 650 m alt., 12. VI.1989, Ignatov 0/669 ( MHA) ; Artyshtu River 0.3 km of its mouth, 600 m alt., 20.IX.1989, Zolotukhin s.n. ( MHA) ; Kukol stow in the valley of Karakem River, 23. VI.1989, Ignatov 0/1055 ( MHA) ; Sebalino District: Katun River valley near Ust-Sema , 380 m alt., 28.VII.1991, Ignatov & Ignatova 24/163 ( MHA) ; Chemal District: Chemal Creek 3 rm upstream mouth, 450 m alt., 12.VII.1993, Ignatov & Ignatova 34/63 ( MHA) ; Irkutsk Province, Irkutsk surroundings, 10.VII.1932, Kostin s.n. ( LE) ; Baikal Lake, Maloe More, Yadyrtui Cape , 19.VII.1987, Bardunov s.n. ( LE) ; Zabaikalsky Territory: Dul’durga District, Alkhanai National Park , Ilya River , 8.VII.2006, Afonina 1606 ( LE) ; Kyra District: Sokhondinsky Biosphere Reserve: Agutsa River , 1098 m alt., 18.VII.2010, Afonina A3710 ( LE) ; Enda River, 1320 m alt., 12.VII.2010, Afonina A1910 ( LE) ; Bukukun River, 1646 m alt., Czernyadjeva 42-11 ( LE) ; Ulota District: Ingoda River ner Leninskiy Settlement , 6.VIII.2011, Afonina 0611 ( LE) ; Krasnochikoisky District: Zhindokon Creek, 963 m alt., 8.VIII.2011, Afonina 1411 ( LE) ; Nerchinsk District: Nerchinsko-Kuenginsky Rangem Marikta River, 15.VII.2012, Czernyadjeva 7-12 ( LE) ; Aleksandrovsky Zavod District, Malaya Borzya River, 817 m alt., 27.VII.2012, Afonina 4412 ( LE) ; Tungokochin District, Ul’durga River, 618 m alt., 16.VII.2012, Afonina 1112 ( LE) ; Chita District, Aratsa River, 964 m alt., 13.VII.2012, Afonina 0312 ( LE) ; Republic of Buryatia: Selenga River near Novoselenginsk Settlement , 4.VII.2007, Afonina 00707 ( LE) ; Amurskaya Province: Selemdzhinsky District: Norskaya Hill, 220 m alt., 21. VI.2011, Bezgodov 391 ( MHA) ; Nora River, 240 m alt., 15.VII.2010, Bezgodov 460 ( MHA) ; Khabarovsk Territory: Verkhnebureinsky District: Chegdomyn , 450 m alt., Ignatov 97-284 ( MHA) ; Sovgavan District: Botchinsky State Reserve: Spokoiny Creek , 290 m alt., 22.VIII.2013, Ignatov & Ignatova 13-1035 ( MHA) ; Mulpa River , 245 m alt., 14.VIII.2013, Ignatov & Ignatova 13-494 ( MHA) ; Primorsky Territory: Partizank District: Chandolaz Mt. , 250 m alt., 8.IX.2013, Ignatov et al. 13-1840 ( MHA) ; Shkotovo District: Ussurijsky Nature Reserve, Koryavaya Pad’ , Zmeinaya Mt. , 331 m alt., 15.X.2008, Ignatov 08-299 ( MHA) ; Republic of Sakha / Yakutia: Khangalassky District : Lenskie Pillars National Park , Lena River downstream Labyja Creek mouth, 150 m alt., 19.VIII.2000, Ignatov 00-164 ( MHA) ; Verkhoyansk District, 30 rm W of Tabalakh Settl., Tuostakh Creek, 5.VIII.2009, Isakova s.n. ( MHA) .

ASIA: GEORGIA: Borzhom, pr. fl. Kura , VIII.1877, A.H. & V.F. Brotherus s.n. ( LE); Imeretia, pr. fl. Rion , 6. VI.1877, A.H. & V.F. Brotherus s.n. ( LE); Ossetia, Balta, ad. fl. Terek , 19. V.1881, A.H. & V.F. Brotherus s.n. ( LE). AZERBAIJAN: Zakatalsky District, Kebeloba Settl. , 14. VI.1969, Lyubarskaya s.n. ( LE); Lenkoran, Bilyasar Mountains , 28. V.1937, Topachevsky s.n. ( LE) .

MONGOLIA: Khentii Province, 33 km NW of Dadal, Bukukun River valley , 4.VIII.2006, Afonina s.n. ( LE) ; Bayan-Hongor Province, Ikh-Bogd Mt. , 2000 m alt., 29. VI.2001, Ignatov 01-479 ( MHA) ; Omnogovi Province, Khuren khanyn khep mountain ridge, 2000 m alt., 25. VI.2001, Ignatov 01-456 ( MHA) .

CHINA: Inner Mongolia, NE of Huhhot, Yingshan Mountains, Manhan Mt. , 1650 m alt., 24.X.2008, Ignatov 08-511 ( MHA) .

EUROPE: AUSTRIA: Carinthia: Drautal, Danielsberg, ca. 900 m alt., 14.VI.2003, Köckinger 15056 ( MW, ex Hb. H. Köckinger); Lavanttal, Waldensteiner Graben E of Twimberg , 700 m alt., 12.VI.2004, Köckinger 15057 ( MW, ex Hb. H. Köckinger); SWITZERLAND: Ticino, Cugnasco-Gerra, Sciarana, 350 m alt., 9.IV.2005, Schnyder 2005070 (ex Herbarium Norbert Schnyder). HUNGARY: Ktyptogamae exsiccatae #1892, col. Lojka ( LE) . ITALY: Südtirol, Meran, no date, Zickendrath s.n. ( LE) .

NORTH AMERICA: U.S.A.: Ohio: Adams Co., Spring Glen National Area , 215 m alt., 22. V.2006, Buck 50414 ( NY) ; Oklahoma: Payne Co., Stillwater Creek, 22.III.1942, Ikenberry 829 ( LE) ; Missouri: Shannon Co., Jam-Up Creek, 28.XI.1961, Redfearn 9484 ( LE) .

Differentiation. Distinctive characters of F. ciliaris include ovate, abruptly acuminate leaves, regularly denticulate margins with small marginal teeth, short laminal cells 20–45 x 18–11 µm with length/width ratio 2– 4:1 and short, ovate capsules. Its difference from F. altaica , F. major , F. pusilla and F. rostrata are discussed in comments to them.

KEY TO IDENTIFICATION OF FABRONIA SPECIES IN RUSSIA

1. Leaves ovate, abruptly long acuminate; leaf length/ width ratio 2.4–2.8:1 ............................................. 2

— Leaves ovate-lanceolate or lanceolate, gradually long acuminate; leaf length/width ratio (2.4–)2.8–3.7:1 .. 4

2. Marginal teeth unequal, large 3–5-celled teeth alter with small unicellular teeth ...................... F. pusilla

— Marginal teeth equal, small, 1(–2)-celled ............. 3

3. Peristome present ..................................... F. ciliaris

— Peristome absent ............................ [ F. matsumurae ]

4(1) Peristome absent; operculum with a curved long beak; leaf margins finely serrulate ......... F. rostrata

— Peristome present; operculum with a short beak or mammillate; leaf margins serrate .......................... 5

5. Upper and middle leaf cells 50–70(–85) µm long, 4– 8:1; teeth at leaf margins 25–50 µm long .... F. major

— Upper and middle leaf cells 35–50 µm long, 4–6:1; teeth at leaf margins 20–30 µm long .......... F. altaica

Figures

Fig. 5. 1–6 – Fabronia ciliaris; 7–10 – F. altaica (group 2); 11 – F.cf. altaica; 12–15 – F. major (group 1). 1 – putative isotype of F. ciliaris, Helvetia, LE; 2 – Russia, Dagestan, Ignatov & Ignatova 09-766, MHA; 3 – Russia, Altai Republic, 3.VII.1989, Ignatov 0/43, MHA; 4 – Russia, Khabarovsk Territory, Ignatov & Ignatova 13-494, MHA; 5 – Amurskaya Province, 21.VI.2011, Bezgodov 391, MHA; 6 – U.S.A., Ohio, Buck 50414, NY; 7 – Russia, Altai Republic, Ignatov & Ignatova 12-747, MHA; 8 – Altai, 31.VII.1991, Ignatov & Ignatova 25/53, MHA; 9 – Russia, Dagestan, Ignatov & Ukrainskaya 09-480, MHA; 10 – Russia, Dagestan, Ukrainskaya 14098, LE; 11 – Italy, 31.X.1898, Artaria s.n., LE; 12 – Austria, 2.X.2013, Köckinger 14991, duplicate in MW; 13 – Austria, 18.IV.2001, Köckinger 14992, duplicate in MW; 14 – Russia, North Ossetia/Alania, Ukrainskaya 15938, LE; 15 – Russia, Dagestan, Ignatov & Ukrainskaya 09-572, MHA. 1–15 – leaves. Scale bar: 0.5 mm for all.