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28BRYOPHYTES — МОХООБРАЗНЫЕ
On the genus Ulota (Orthotrichaceae, Bryophyta) in Russia
V. E. Fedosov, E. A. Ignatova
Lomonosov Moscow State University, Faculty of Biology, Moscow Corresponding author. V. E. Fedosov, [email protected]
Abstract. Specimens of the genus Ulota kept in the bryological herbaria of MW, MHA, LE, NSK, IRK, UUH, VLA and VBGI were revisited basing on the recent taxonomical treatments of the genus in Europe, Asia and North America. Among ten species of the genus previously known from the territory of Russia, U. phyllantha was recently transferred to the new genus, Plenogemma, while Ulota crispa, which has been considered before in the broad sense, was proved to include at least three separate species, U. crispa s. str., U. intermedia and U. crispula; all three species occur in Russia. According to our results, diversity of the genus within North East Asian center (cf. Garilleti et al., 2015) was underestimated. Two new species from the of U. japonica affinity, U. orientalis sp. nov. and U. pacifica sp. nov. are described herein from north Pacific Region (Khabarovsk Territory and South Kurils correspondingly). Thus, at least thirteen taxa of the genus are presently known in Russia; distribution data were significantly corrected for some of them. Key to identification of Ulota species in Russia and adjacent areas is provided; their distribution is discussed and mapped
Keywords: Ulota orientalis, Ulota pacifica, biogeography, moss flora, taxonomy, Asia, Russia.
О роде Ulota (Orthotrichaceae, Bryophyta) в России
В. Э. Федосов, E. A. Игнатова
Московский государственный университет имени М. В. Ломоносова, Биологический факультет, Москва, Россия
Автор для переписки: В. Э. Федосов, [email protected]
Резюме. Проведена ревизия образцов рода Ulota, хранящихся в бриологических гербариях MW, MHA, LE, NSK, IRK, UUH, VLA и VBGI на основании современных таксономических обработок рода для Европы, Азии и Северной Америки. Из десяти видов, приводившихся ранее для России, U. phyllantha в последнее время была перенесена в монотипный род Plenogemma, а для Ulota crispa показана применимость более узкой концепции видов с выделением U. intermedia и U. crispula; все три вида встречаются в России. Разнообразие северо-восточноазиатского центра диверсификации рода (cf. Garilleti et al., 2015) оказалось недооцененным: из Северной Пацифики (Хабаровский край и Северные Курилы соответственно) выявлены и описаны два новых вида родства U. japonica - U. orientalis sp. nov. и U. pacifica sp. nov. Таким образом, в России встречается не менее 13 видов рода, и представление о распространении некоторых из них существенно изменились; обсуждается их распространение, приводится ключ для определения видов рода Ulota флоры России.
Ключевые слова: Ulota orientalis, Ulota pacifica, биогеография, бриофлора, татеономия, Азия, Россия.
Introduction
At the moment, the number of accepted species in the genus Ulota D. Mohr varies from 64 (Tropicos, 2017) to 69 (Caparros et al., 2011, Garilleti et al., 2015); they are widespread in temperate climates of both Hemispheres mostly under oceanic to hyperoce-anic climatic conditions. Garilleti et al. (2015) underlined three diversity centers of the genus within Palearctic, i.e., Europe (11 species), Altai Mts. (4 species) and Pacific region south from the Kamchatka Peninsula (13 species), while in the remaining area only one species, U. curvifolia (Wahlenb.) Sw. occurs. According to Ignatov et al. (2006), ten species of the genus were previously known in Russia. However, in last decade a lot of taxo-nomical issues within the genus have been improved or clarified, causing a need of circumstantial revision of the genus in Russia.
Taxonomical revision of the genus Ulota in Asia (Wang, Jia, 2012) provide a suitable keys and descriptions for the examination of specimens and some recent data about the species distribution, though data on Russian Ulota specimens are omitted and species distribution in Russia is only briefly mentioned. In particular, Wang and Jia (2012) consider U. reptans Mitt. as a Japanese endemic, thus challenging the identification of Russian specimens reflected in Igna-tov et al., 2006. Further study of Ulota in Japan (Suzuki, Iwatsuki, 2013) revealed some new species for East Asian biodiversity center and resurrected specific level for U. longifolia Dixon et Sakurai, which was considered to be conspecific with U. crispa (Hedw.) Brid. by Noguchi and Iwatsuki (1989). Due to dioicous sexual condition and numerous gemmae at upper leaf apices, U. phyllantha Brid. was segregated in a separate genus Plenogemma Plasek, Sawicki et Ochyra (cf. Plasek et al., 2015). Caparros (2015) provided some essential comments on morphology of some poorly understood Ulota species. Finally, recent integrative taxonomical study (Caparros et al., 2016) resulted in resurrection of a species status for U. intermedia Schimp. and U. crispula Bruch, which for a long time have been considered to be conspecific with U. crispa. Authors provided a key to species identification and data on their world distribution, where all three species were reported from Russia though based on single or few specimens. On the other hand, recent field works in southern Far East earned numerous Ulota collections, which needed critical study in light of recent advances in taxonomy of this genus.
We revised the specimens of the genus Ulota kept in bryological herbaria MW, MHA, LE, NSK, IRK, UUH, VLA and VBGI using the morphological conception of species provided by recent taxonomical treatments (Ignatov, Ochyra, 1994; Garilleti et al., 2000; Smith, 2004; Lonnell, 2008; Wang, Jia, 2012; Vitt, 2014; Caparros et al, 2014, 2016; Caparros, 2015; Blockeel, 2017).
Our results proved the occurrence of all species listed in Ignatov et al. (2006) in Russia as well as all-three species of U. crispa complex considered by Caparros et al. (2016).
Furthermore, several specimens of the Ulota japonica (Sull. et Lesq.) Mitt. affinity from Russian Far East do not fit to the morphological conception of any of known species. Unique specimen from the coastal area of sea of Okhotsk combines characters of U. japo-
nica and U. rehmannii Juratzka, though differs from both due to differentiated perichaetial leaves and calyptrae with numerous long hairs. Two specimens from south Kuril Islands share gametophyte characters of U. japonica, but have short, curved setae, weakly ribbed, cylindrical, slightly inflated, not contracted below the mouth capsules and narrow, filiform segments. In addition, the specimens from Iturup Island were collected on the coastal rock, which is uncommon habitat for this mostly corticolous genus, while in the specimen from Shikotan Island the plants described above were intermixed with true U. japonica, demonstrating contrasting morphology despite growing in the same environments. Thus, in our opinion, these entities are morphologically distinct and deserve to be considered as a separate species. Their formal description along with introducing new data on the other species distribution is a matter of a present paper.
Ulota orientalis Fedosov et Ignatova sp. nov. (Plates I, II, Fig. 1)
Type: Russia, Khabarovsk Territory, 26 VIII 2013, Ignatov, Ignatova 13-1218 (Holotype MHA!, Isotype MW!).
Diagnosis: The species being described combines thin-walled rectangular proximal leaf cells, well differentiated perichaetial leaves and rather small spores, characteristic for U. barclayi Mitt. and turbinate, not contracted below mouth capsules with split pairs of exo-stome teeth strolate on distal OPL surface and narrow endostome segments, characteristic for U. rehmannii and U. megalospora Venturi; in addition, unlike in both groups, calyptra in U. orientalis is covered throughout by rather long straight hairs.
Etymology: The species name is given due to occurrence in the Far East (Asia orientalis).
Description. Plants in dense tufts, green to brownish downward; stems erect or ascending, not creeping, sparsely ramified, 1-2 cm long, in basal portion weakly tomentose. Leaves slightly curved when dry, spreading when wet, 1.6-2.4 x 0.4-0.6 mm, lanceolate from a slightly widened, concave base, acute to acuminate; upper leaves gradually becoming larger, 2.4-3.0 x 0.7-0.9 mm, short lanceolate from ovate base; margins weakly recurved in middle part of leaf on one side or plane, in upper part papillose-crenulate; costa vanishing few cells below apex or, in lower acuminate leaves, subpercurrent, brownish proximally, smooth on abaxial and adaxial sides; basal laminal cells near costa (17-)19-36(-40) x 9-14 ^m, thin- to moderately thick-walled, rectangular to elongate-rectangular, longitudinal cell walls not or slightly sinuose, often with slightly oblique, thickened transverse walls, basal marginal cells quadrate to short rectangular, differentiated in 8-13 rows, 10-18 x 9-14 ^m, thin-walled or with slightly thickened transverse walls, pellucid, or all basal leaf portion colorless, with rather sharp gently V-shaped transition to upper leaf cells, smooth; upper leaf cells rounded-hexagonal, rounded-rhomboidal, transverse-elliptic (predominantly in middle leaf portion), subquadrate to oblong (predominantly in upper leaf portion), (6-)8-12(-14) x 8-12 ^m, moderately thick-walled, chlorophyllous, with few
Fig. 1. Distribution of Ulota orientalis (star), U. pacifica (hollow triangle), U. bruchii (hollow circle) and U. coarctata (solid triangle) in Russia.
scattered low simple papillae, no more than one per cell. Perichaetial leaves differ from stem leaves mostly in larger size, rounded tips and in having broadened base, thus their shape is ovate to ovate-lanceolate; leaf tips obtuse to round. Seta 1.8-2.4 mm. Capsule mostly turbinate when mature, 1.6-2.4 mm, not contracted at mouth, evenly tapering to seta, brown, with 8 broad longitudinal ribs ca. % to entire capsule length, composed of 5-6 cell rows and with narrow furrows between them; exothecial bands are differentiated neither in coloration nor in cell pattern, since whole exothecium is composed of nearly equally broad cells with incrassate longitudinal walls and thin transverse walls; stomata in neck and proximal portion of capsule, reaching capsule middle; peristome double; exostome teeth arranged pairwise but not fused, narrowly lanceolate, 380-510 ^m, reflexed, straight, adherent to the outer capsule wall, pale orange; outer peristomial layer (OPL) strongly papillose proximally, weakly papillose, covered with longitudinal ridges in distal third; primary peristomial layer (PPL) with low elongate papillae proximally, papillose distally; 1-2 uppermost tooth cells often weakly fenestrate; endostome segments 8, about half of teeth length, broadly linear, composed of one row of cells, incurved, slightly papillose. Operculum not seen. Calyptra narrow-conic, with long, straight, upward directed hairs. Spores spherical, 18-22 ^m, green brownish, papillose.
Differentiation. Rather thin-walled rectangular basal leaf cells near costa with neither sinuose nor porose longitudinal cell walls in combination with small spores differ Ulota orientalis from most of Holarctic Ulota species excepting U. japonica, U. pacifica (see below) and north American U. barclayi Mitt. On the other hand, turbinate capsules, gradually narrowed from the mouth to the seta and striolate distal OPL surface remind rather U. megalospora and U. rehmannii than these species. 144
Ulota barclayi, the only previously known Pacific Ulota species with differentiated perichaetial leaves, has been recently considered to be conspecific with U. japonica (cf. Anderson et al., 1990). Despite Vitt (2014) considered U. barclayi as a separate species, in comments he underlined that in fact it is closely related and probably conspecific with U. japonica. According to his treatment, the only difference between them includes strongly differentiated perichaetial leaves in U. barclayi vs. not differentiated in U. japonica; however, according to the provided key, U. barclayi can have weakly differentiated perichaetial leaves as well. The description provided by Vitt (2014) fits both species in other respects and thus seems to be broadened to accommodate the specimens that we would prefer to place in U. japonica. According to Caparros (2015) and Lara (pers. com.), U. barclayi differs from U. japonica in having angulate, red rimmed opercula, which are circular, not red rimmed in the latter species.
From U. japonica, U. orientalis differs in having obtuse to rounded (vs. acute) perichaetial leaves; oblong to funnel-shaped, not constricted below mouth, brown (vs. mostly urceolate, markedly constricted below mouth, yellowish to light brownish) capsules; weakly differentiated (vs. well differentiated) exothecial bands; much longer, not fused (vs. shorter, mostly tightly fused) exostome teeth and slender, not keeled endostome segments, composed of one cell row (vs. more robust, keeled, composed of two cell rows). Differentiation of U. orientalis and U. pacifica is considered under this species.
From somewhat similar habitually U. rehmannii, U. orientalis differs in smaller spores, weakly incrassate basal leaf cells, perichaetial leaves with obtuse to rounded apices, and brown (vs. pale brownish) capsules.
Distribution and ecology. Corticolous on alder trunk in alder crooked forest; known from a single locality on the seashore of the Gulf of Tatary (48°58'17.8"N, 140°22'18.7"E, 120 m. alt.) in Sovetskaja Gavan' town outskirts, Tsvetuschaja Polyana area.
Ulota pacifica Fedosov et Ignatova sp. nov. (Plates III, IV, Fig. 1)
Type: Russia, Sakhalin Region, Iturup Island 11 IX 2015, Fedosov 15-2-154 (Holo-type MW!, Isotype MHA!).
Diagnosis: The species being described superficially resembles U. japonica due to thin-walled rectangular proximal leaf cells and rather small spores, but differs from it in having short, curved setae, weakly ribbed, not contracted below mouth cylindrical capsules with poorly differentiated exothecial bands and narrow, filiform segments.
Ethymology: The species name is given due to its occurrence in Pacific region.
Description. Plants in dense tufts, yellowish-green or dark green; stems erect or ascending, sparsely branched, 0.5-1 cm long, in basal portion moderately tomentose. Leaves slightly curved when dry, spreading when wet, 1.5-2.5(2.8) x 0.4-0.7 mm, lanceolate from a slightly widened, concave base, acute to acuminate; margins plane or weakly recurved in lower part of leaf on one side, entire; costa vanishing few cells below apex or subpercurrent, smooth on abaxial and adaxial sides; basal laminal cells near costa 35-60 x 8-10 ^m, elon-
gate-rectangular, with thin or moderately and slightly unevenly thickened longitudinal walls, basal marginal cells quadrate to short rectangular, differentiated in 6-7 rows, 12-22 x 1012 jim, with thin longitudinal walls and strongly thickened transverse walls, pellucid, gradually transiting to upper leaf cells; upper and median leaf cells subquadrate, rectangular and rhomboidal, 8-14 (-18) x 8-10 jm, moderately thick-walled, chlorophyllous, leaf lamina smooth or with few scattered low simple papillae. Perichaetial leaves not differentiated from stem leaves, ca. 2.5 mm long and 0.8 mm wide at base. Seta 0.9-1.2 mm, arcuate. Capsule with oblong-ovate urn 1.2-1.4 mm long and 0.8-0.9 mm wide, abruptly narrowed to the neck 0.8-0.9 mm long, urn not contracted below the mouth after spore release, pale brownish, with 8 broad longitudinal ribs ca. 3% of urn length, composed of 5-6 cell rows and with wide, shallow furrows between them; exothecial bands not differentiated in color, composed of larger cells with thicker walls; stomata in neck and proximal portion of urn; peristome double; exostome teeth narrowly lanceolate, brownish, fused in pairs, becoming slightly cleft in upper part with age, 270-290 jm, reflexed, straight, adherent to the outer capsule wall, pale yellow; outer surface (OPL) densely papillose in distalmost plates, with oblique ridges and dense papillae in the most part of teeth; endostome segments 8, 190-205 jm long, filiform, composed of one row of cells, widened in basalmost part, incurved, slightly papillose on inner surface. Calyptra conic, with scattered long, straight, upward directed hairs. Spores spherical, 17-21 jm, green brownish, papillose.
Differentiation: Ulota pacifica differs from the most closely related U. japonica in curved vs. straight setae; capsules cylindrical, slightly inflated to ovate, not contracted below mouth, weakly ribbed to nearly smooth vs. mostly urceolate, contracted below mouth, strongly ribbed; endostome segments narrow, filiform, widened proximally only in basalmost part vs. broader, linear, gradually widened proximally; among gametophyte characters, smooth leaf lamina or with few scattered papillae of U. pacifica should be mentioned; it differs from regularly papillose distal leaf lamina of U.japonica. The same characters and not differentiated perichaetial leaves differ it from U. barclayi. Ulota orientalis has quite remarkable differentiated perichaetial leaves and turbinate capsules. In having rather short, often curved setae and saxicolous growth, U. pacifica resembles U. curvifolia, but differs from the latter in thin-walled vs. thick-walled proximal leaf cells; absent or few, single vs. regularly present, mostly forked papillae in distal leaf cells; never urceolate vs. more or less urceolate mature capsules; weakly vs. strongly differentiated exothecial bands and papillose vs. ridged distally exostome teeth. East Asian U. perbreviseta Dixon et Sakurai resembles U. pacifica in having short curved setae, capsules not constricted below mouth and narrow endostome segments, but differs in having crisped leaves and thick-walled proximal leaf cells.
Distribution and ecology. Corticolous or saxicolous. This species is known from two localities in southern Kuril Islands: in northern part of Iturup Island, faced to Pacific Ocean, shore on Tsirk Bay, (45°19'47.5"N, 148°37'10.4"E, 10 m. alt.), on coastal vol-canogenic cliff of neutral composition, and in Shikotan Island near Malokurilsk Settlement
Plate I. Ulota orientalis. 1 — capsule; 2 — habit, dry; 3 — calyptra; 4 — peristome and exothecium;
5 — leaf transverse section; 6 — distal leaf cells; 7 — median leaf cells; 8-9, 11 — stem leaves; 10 — perichaetial leaf; 12 — basal leaf cells. Scale bars: 2-3 — 2 mm, 8-11 —1 mm, 1 — 0.5 mm 4 — 200 pm, 5-7, 12 — 100 pm. All from MHA 13-1218.
Plate II. Ulota orientalis. 1 — peristome; 2 — basal tooth portion from the inside (PPL); 3 — basal tooth portion from the outside (OPL); 4 — spore; 5 — distal tooth portion from the inside (PPL); 6 — distal tooth portion from the outside (OPL); 7 — endostome segment. Scale bars: 1 — 100 pm, 2-4 — 5 pm, 5-7 — 10 pm. All from MHA 13-1218.
Plate III. Ulota pacifica. 1, 5 — capsules; 2 — habit, dry; 3 — leaf transverse section; 4 — calyptra; 6-8, 13 — stem leaves; 9 — distal leaf cells; 10 — median leaf cells; 11 — peristome and exothecium; 12 — endostome segment; 14 — basal leaf cells. Scale bars: 1-2 — 2 mm, 6-8, 13 — 1 mm, 4-5 — 0.5 mm, 11-12 — 200 pm, 9-10, 14 — 100 pm. All from MW 15-2-154.
Plate IV. Ulota japonica from MHA K-36-33-07 (1, 3) and U. pacifica from MW 15-2-154 (2, 4). 1-2 — peristome; 3-4 — inner surface of peristome teeth (OPL). Scale bars: 1-2 — 100 pm, 3-4 — 10 pm.
(ca. 43°52'N, 146°51'E, 100 m. alt.), in wet brook hollow with alder thickets. Apparently, U. pacifica occurs in Japan as well, but so far it has not been segregated from U. japonica. At least, in the image provided by Wang et Jia (2012), the plant illustrated in Fig. 4D has the same urceolate capsules as in Russian specimens, while the plant illustrated in Fig. 4E with its ovate capsule, in our opinion, better corresponds to U. pacifica.
Other specimens examined: Russia, Sakhalin Region, 23 VIII 2007, Bakalin, MHA.
Basing on the recent publications cited above, we compiled the key to the species of Ulota in Russia and adjacent areas, which has later been updated in the course of identification of the Russian specimens. The provisional species in the key are placed in brackets.
Key to the species of Ulota in Russia and adjacent areas
1. Capsules fusiform to pyriform, distinctly narrowed toward small mouth......................................2
— Capsules rounded, oblong, cylindrical, urceolate or funnel-shaped, rarely pyriform, often strongly contracted below mouth, but usually not at mouth.....................................................................4
2. Mature capsules when and after spore releasing inflated, mostly smooth, with weak ribs only just below mouth................................................................................................................U. coarctata
— Mature capsules when and after spore releasing with longitudinal ribs on most of their length .... 3
3. Leaves curved when dry; peristome simple.............................................................U. drummondii
— Leaves crisped when dry, peristome double.....................................................................U. bruchii
4(1). Mostly saxicolous species.............................................................................................................5
— Mostly corticolous species.............................................................................................................7
5. Leaves straight to curved when dry, plants growing in cushions; papillae simple.........................6
— Leaves contorted when dry, plants growing in creeping patches; papillae partly forked .....................................................................................................................................U. curvifolia
6. Setae very long, 3-4 mm, straight; mostly European species...................................U. hutchinsiae
— Setae short, 1.5-2 mm, mostly curved; East Asian species............................................U. pacifica
7(4). Leaves strongly contorted or crisped when dry; calyptra densely hairy........................................8
— Leaves curved to contorted when dry, calyptra mostly with few hairs to naked..........................11
8. Leaves contorted to slightly crisped when dry, from scarcely widened, weakly concave elliptic base rather gradually narrowed to acumen; basal marginal cells weakly differentiated, in 1-3 rows; endostome segments uniseriate or with biseriate patches, cells thin-walled.........U. crispula
— Leaves strongly crisped when dry, from wide, concave base abruptly narrowed to acumen; basal marginal cells sharply differentiated, in (2)3-9 rows; endostome segments uniseriate, with thick transverse cell walls.......................................................................................................................9
9. Mature capsules very narrow, when dry and empty strongly constricted below mouth, with robust orange ribs and narrow furrows between them; exothecial bands composed of 4-6 cell rows..................U. crispa
— Mature capsules wider, cylindrical, when dry and empty not or weakly constricted below mouth, with less prominent yellowish ribs, composed of 2-4(5) cell rows, and rather wide light furrows between them .............................................................................................................................. 10
10. Capsules with reddish rim at mouth; teeth reddish, bordered with hyaline halo; endostome segments 16, filiform; spores 15-20(23) pm..................................................................[U. longifolia]
— Capsules not rimmed; teeth pale, without hyaline halo; endostome segments mostly 8, broadly linear; spores mostly 18-35 pm.................................................................................U. intermedia
11(7). Basal leaf cells near costa rectangular, with thin to moderately thick walls, longitudinal cell walls straight or weakly sinuose; spores 15-24 pm .............................................................................12
— Basal leaf cells near costa elongate, strongly thick-walled, with more or less sinuose and porose longitudinal walls; spores 23-40(60) pm.....................................................................................15
12. Perichaetial leaves differentiated, obtuse to rounded; capsules not contracted below mouth, funnel-shaped, rarer oblong when mature, brown; exostome teeth 16, not fused in pairs, 380-510 pm.................................................................................................................U. orientalis
— Perichaetial leaves weakly or not differentiated, often somewhat larger than lower leaves, acute to acuminate; capsules urceolate, cylindrical to ovate, mostly contracted below mouth, yellow to brownish; exostome teeth mostly fused in 8 pairs, up to 350 pm ...............................................13
13. Setae mostly curved, capsules cylindrical, slightly inflated to ovate, not contracted below mouth, weakly longitudinally ribbed; segments narrow, filiform...............................................U. pacifica
— Setae straight; capsules urceolate, not inflated, contracted below mouth, strongly longitudinally ribbed; segments wider, linear ....................................................................................................14
14. Perichaetial leaves mostly differentiated; operculum red-rimmed, angulate...............[U. barclayi]
— Perichaetial leaves not differentiated; operculum rounded, not rimmed .......................U. japonica
15(11). Leaves mostly with acuminate apices; stomata in middle portion of capsules; spores
35-60 pm.............................................................................................................[U. megalospora]
— Leaves with acute apices; stomata in proximal portion of capsules; spores 21-40 pm................16
16. Leaves (1.4)1.5-2.4 mm long; upper and median laminal cells weakly papillose, papillae low;
basal marginal cells differentiated in 3-7 cells rows; exostome teeth in upper portion with lines (striae) and papillae, to nearly smooth........................................................................U. rehmannii
— Leaves 1.0-1.5 mm long; upper and median laminal cells clearly papillose, papillae high; basal marginal cells differentiated in 1-3 cells rows; exostome teeth not or weakly striolate, papillose throughout........................................................................................................................U. reptans
Since the full taxonomic treatment itself is to be published separately in the fourth volume of «The moss flora of Russia», here we present only brief differentiation of the remaining Ulota species previously known in Russia and actual data on their distribution.
Ulota bruchii Hornsch. ex Brid., 1827, Bryol. Univ. 1: 794. (Fig. 1)
Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 57.
Differentiation. According to Garilleti et al. (2000), the most useful character to separate O. bruchii from the species of U. crispa affinity is the longitudinally striolate en-dostome segments. Its fusiform capsules narrowed to the mouth differ a little from capsules of U. intermedia and U. crispula, which are cylindrical, while U. crispa s. str. has capsules strongly constricted below mouth. Contrary to U. intermedia and U. crispula, exothecial bands with thick longitudinal walls reach the urn mouth in U. bruchii; in this character it is similar to U. crispa, but it has contrastingly different shape of capsules after spore release.
Distribution and ecology. An exclusively European distribution of Ulota bruchii seems to be very sensitive to the moisture condition. Even in Britain, where the species is widespread, it occurs mostly in more humid conditions (Smith, Hill, 1975). According to Blockeel et al. (2014), in Britain and Ireland these two species may occur together, but U. bruchii is commoner than U. crispa in many districts, while U. crispa is more frequent in upland areas than U. bruchii. It is rather frequent in the Alps and in SW Norway. Few collections of the species are known from Portugal, Spain, France, Belgium, Netherlands, Denmark, North Germany, Slovakia, Finland, and Baltic countries. In Russia the species
occurs in eastern extremity of its distribution in maritime areas of the Kaliningrad Region (Curonian Spit) and in the Leningrad Region (Islands of Baltic Sea). Exclusively cortico-lous, usually on aspen.
Selected representative specimens: Russia, Leningrad Region, 3 VII 2006, Kurbatova, LE.
Ulota coarctata (P. Beauv.) Hammar, 1852, Monogr. Orthotrich. Ulot. Suec. 25. (Fig. 1) Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 58.
Differentiation. The species is quite remarkable due to inflated, pyriform, nearly smooth mature capsules, abruptly narrowed toward the star-like mouth; its differentiation is obvious and is reflected in the key above. Among other characteristic features, rather long acuminate, curved leaves along with not fused, rather short exostome teeth and mostly lacking endostome segments should be mentioned.
Distribution and ecology. Ulota coarctata is a nicely pronounced amphiatlantic cool temperate/hemiboreal species, which is rather frequent in Britain, SW Scandinavia and Quebec, with scattered localities in Western Europe, Alps, Germany, Tatras, Ontario and Eastern part USA (Smith, 2004; Lonnell, 2008; Ros et al, 2013; Plasek et al, 2016; Vitt, 2014; etc). Eastern extreme of its distribution is in the Caucasus (Caucasian and Teberda state reserves). Corticolous, mostly on alder; suboceanic, very sensitive to the moisture condition.
Selected representative specimens: Russia, Krasnodar Territory, 20 VI 2000, Akatova, MHA.
Ulota crispa (Hedw.) Brid., 1819 [1818], Muscol. Recent. Suppl. 4: 112. (Fig. 2)
Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 59.
Differentiation. According to Caparros et al. (2016), the species differs from closely related U. crispula and U. intermedia by urceolate capsules, which are strongly contracted below mouth, wide orange to reddish ribs, narrow furrows, collapsing at capsule constriction, exostome teeth, which are not split even in old dry capsules, with hyaline halo along margins. Most of these features excepting capsule shape are characteristic to the Japanese endemic U. longifolia, which until present has been considered as subspecies of U. crispa. It differs from U. crispa s. str. in having cylindrical, not or slightly contracted below mouth capsules with light ribs, composed of elongate cells, longer peristome teeth (325-360 pm), and 16 filiform endostome segments vs. mostly 8 (occasionally with 8 additional) wide linear segments, thus combining characters of two species, U. intermedia and U. crispa. Despite the special attention, no one specimen with character combination fitting U. longifolia was found in the Russian material. From the other Ulota species U. crispa differs in having wide, concave leaf base, abruptly tapering to distal leaf lamina, and strongly crisped leaves.
Distribution and ecology. According to Caparros et al. (2016), U. crispa has temperate amphioceanic distribution. It is widespread in the South and Middle Europe, penetrating northward to southern Norway and eastward to Turkey, Caucasus and Transcarpathia. In East Asia the species occurs in China, Japan, and Taiwan; in North America it is present along Pacific
Fig. 2. Distribution of Ulota crispa (hollow circle) and and U. drummondii (solid circle) in Russia.
coast, mostly in British Columbia and Washington with few inland localities, but seems to be absent in Atlantic North America. In Russia the species occurs in western part of Russian Caucasus, there it is especially abundant in southernmost part of Black Sea coastal area, Khosta Taxus and Buxus dominated forest (see Ignatov et al., 2002) eastward to Kabardino-Balkarian Republic; few specimens are known from Primorye Territory in the Russian Far East. All specimens from the middle part of European Russia, Altai and other parts of the Russian Far East previously identified as U. crispa were referred to U. intermedia (cf. Caparros et al., 2016). Corticolous.
Selected representative specimens: Russia, Krasnodar Territory, 9 IX 2008, Seregin, Ashurkova, Vinokurova, MHA; Karachayevo-Circassian Republic, 16 IX 2005, Ignatov, Ignatova, MHA.
Ulota crispula Bruch, 1827, Bryol. Univ. 1: 793. (Fig. 3)
Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 60.
Differentiation. This species differs from U. crispa and U. intermedia by curved to weakly crispate leaves, weakly differentiated, narrow group of cells along basal leaf margin, exothecial bands consisted of 2-4 rows of evenly yellowish cells and partly biseriate endostome segments (see Caparros et al., 2016 and the key above). Due to weakly widened leaf base and contorted rather than crisped leaves it could be confused with the other Ulota species not related to U. crispa. From Ulota japonica, U. crispula differs in the pattern of basal leaf cells and mostly larger spores, from U. barclayi, U. rehmannii and U. reptans, in capsules slightly contracted below mouth after spore release and weakly differentiated basal marginal group of cells. Additional comments on the differentiation between U. crispula and U. intermedia see under the latter species. Ulota bruchii differs
Fig. 3. Distribution of Ulota crispula s. str. (soild circle) and putatively closely related Asian taxon (hollow circle) in Russia.
from U. crispula in having capsules distinctly tapered toward the small mouth, which is star-shaped when viewed from above and in longitudinally striolate endostome segments. In Asian part of Russia somewhat problematic specimens, which are hard to be referred either to U. intermedia or to U. crispula occur. In gametophyte characters (rather strongly crispate leaves, shape of leaf base, well differentiated rather broad band of cells along basal leaf margin) they fit to the first species, most of the sporophyte characters (yellowish throughout cells of exothecial bands, fragile, partly biseriate endostome segments) are characteristic for U. crispula, while capsules notably contracted below mouth fit better to U. crispa. Specimens of this kind are most widespread on the Kamchatka Peninsula, while pure U. intermedia is rare in this area and pure U. crispula apparently does not occur in Asia (see Fig. 3), like it was mapped by Caparros et al., 2016. Status of this Asian taxon, its relation to U. crispula and U. intermedia needs to be clarified in the course of overall revision of U. crispa complex in Pacific region, which is quite actual.
Distribution and ecology. The species has cool temperate/hemiboreal, moderately oceanic distribution. Caparros et al. (2016) considered U. crispula as an amphiatlantic species, penetrating to European Russia in eastern limit of its European range. Since for a long time it has not been separated from U. crispa, its distribution is still insufficiently known; our data provide previously unknown Asian extension in the species range. In Russia this species is rare and occurs on eastern edge of its mostly European distribution. Caparros et al. (2016) documented it from two localities in the western Caucasus, in the Black Sea coastal area and in Kabardino-Balkaria, but they also proposed rather wide distribution of the species in lowland European Russia, which was not confirmed by our data. U. crispula was
also found in Teberda State Reserve (Karachayevo-Circassian Republic), North Ossetia, Altai Mountains, Khabarovsk Territory (Botchi State Reserve), Kamchatka Peninsula (and it is rather common there), Sakhalin and Southern Kuril Islands (Fig. 4). Corticolous.
Selected representative specimens: Russia, Karachayevo-Circassian Republic, 19 VIII 1955, Abramov, Abramova, LE; Kabardino-Balkarian Republic, 23 VIII 2005, Ignatov, Ignatova, Kharzinov, MHA; Republic of Altai, 11 VII 1991, Ignatov, MHA; Kamchatka Territory, 22 VII 2004, Czernyadjeva, LE.
Ulota curvifolia (Wahlenb.) Lilj., 1816, Utkast Sv. Fl. (ed. 3): 546. (Fig. 4)
Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 61.
Differentiation. Saxicolous growth in shady niches on overhanging rock surfaces, contorted leaves, spherical to short urceolate capsules, densely hairy calyptrae and exo-stome teeth longitudinally striolate above are quite remarkable to separate U. curvifolia from all other Holarctic Ulota species.
Distribution and ecology. Boreal montane species. Compared with all other Ulota species, U. curvifolia has mostly more northern distribution, or occurs in higher altitudinal belts (Ignatov, Ochyra, 1994; Lonnell, 2008; Wang, Jia, 2012; Vitt, 2014, etc.). With its circumholarctic distribution, in Europe (outside Russia) U. curvifolia occurs only in Scandinavia and on islands of the Baltic Sea; in America it is distributed throughout Canada (but does not occur in USA excepting Alaska); it is the only Ulota species known in Greenland; in Asia outside Russia it is known from a single locality in Chinese Altai. In Russia it is rather widespread, being the only representative of the genus in subarctic and continental mountain areas wherever acidic crystalline bedrocks occur, but it is still unknown in the high Arctic mountains composed of basic to neutral rocks (Putorana Plateau),
schists (Verkhoyansky Range) and volcanic areas (Kamchatka). It also seems to absent in temperate mountain systems, since despite of numerous records of the species in Caucasus and Middle Asia (Manakyan, 1995; Mamatkulov et al., 1998; Kharzinov et al., 2004), we did not find any specimens from those areas excepting the record from Kazakh part of Altai Mts.; in Mongolia it sporadically occurs in highland areas with permafrost occurrence; it is still unknown in Sikhote-Alin Range. In European Russia U. curvifolia occurs in its north-western part where rocks of Baltic Crystalline Shield outcrops are widespread, from the Murmansk Region to western part of the Leningrad Region, in the Ural Mts. — from the Polar Urals to the Bashkortostan Republic, in Altai Mts., West and East Sayans, Anabar Plateau, montane areas around Baikal Lake, Transbaikalia, southern part of Yakutia, the Magadan Region, continental and Beringian Chukotka, and continental part of the Khabarovsk Territory (Fig. 4).
Saxicolous. According to our observation, Ulota curvifolia is especially abundant in granitic and gneiss rocks, where it occupies shaded overhanging surfaces and usually forms extensive mats; sometimes it also occurs on meta-gabbro.
Selected representative specimens: Russia, Murmansk Region, 5 VIII 1948, Schljakov, LE; Republic of Karelia, 6 VII 1975, Volkova, LE; Leningrad Region, 19 VII 2004, Kurbatova, LE; Tyumen Region, 30 VIII 1956, Kildyushevsky, LE; Republic of Altai, 27 VI 1991, Ignatov, MHA; Irkutsk Region, 3 VII 1984, Bardunov, LE; Republic of Buryatia, 11 VII 2008, Afonina, Tubanova, LE; Trans-Baikal Territory, 3 VII 2013, Afonina, LE; Republic of Sakha (Yakutia), 10 VII 1991, Ivanova, LE; Chukotka Autonomous Area, 9 VII 1979, Afonina, LE; Magadan Region, 24 VIII 1976, Blagodatskikh, LE; Amur Region, 11 VIII 1980, Abramova, MW; Khabarovsk Territory, 24 VII 1992, Borisov, MW.
Ulota drummondii (Hook. et Grev.) Brid., 1826, Bryol. Univ. 1(1): 299. (Fig. 2)
Representative illustration: Ignatov et al, 2018 (in prep.): Fig. 62.
Differentiation. This species can be easily distinguished even in the field by combination of the following features: weakly curved leaves; capsules gradually narrowed to the mouth and not contracted below it, strongly furrowed in upper part, causing irregularly angular to star-like shape of the capsule mouth; and simple whitish peristome composed of teeth directed upward to spreading (fragile, often early broken off). However, two East Asian Ulota species, U. gymnostoma S. L. Guo, Enroth et Virtanen and U. delicata Q. H. Wang et Y. Jia should be taken into account since they have somewhat similar sporophyte characters. According to Wang and Jia (2012) they differ from U. drummondii in lacking (U. gymnostoma) or double (U. delicata) peristomes and longer, lanceolate to linear-lanceolate leaves, which are much stronger contorted to crispate when dry vs. mostly ovate lanceolate, curved to slightly contorted in U. drummondii. Differentiation among U. drum-mondii, U. coarctata and U. bruchii are provided in the key above.
Distribution and ecology. This species is characterized by amphioceanic distribution in regions with cool temperate to boreal hyperoceanic climate. It is widespread and
common in wet areas of northern Europe. For example, it is the most common Ulota species in coastal area and along the fjords of SW Norway (Lonnell, 2008; authors' observation), but is known only from few localities southward in Saaremaa Islands and continental Estonia (GBIF, 2017b); in Britain it is common in Scotland (cf. Smith, 2004, Blockeel et al, 2014) and declines southward, with few records in Ireland; it was also reported from the north of Spain (Caparros, 2015). In North America, according to Vitt (2014), the species is also strictly confined to a narrow area of the Atlantic coast (Newfoundland and Acadia) and the Pacific coast (from Washington to South Alaska and along Aleutians). In Asia U. drummondii occurs in Japan and the Russian Far East. Distribution of the species in Russia is generally associated with the insular part of southern Russian Far East, i.e., Sakhalin and Kuril Islands, where it is the most common species of the family. It is also rather frequent in western and southern Kamchatka; few localities are known from the coastal areas of Primorye and Khabarovsk Territories. In European Russia it occurs on islands of the Baltic Sea, mostly inhabits barks and branches of willow, poplar and other deciduous trees, often with other Ulota species and Lewinskya sordida (Sull. et Lesq.) F. Lara, Garilleti et Goffinet.
Selected representative specimens: Russia, Leningrad Region, 8 VIII 2005, Leushina, LE; Kamchatka Territory, 6 VIII 2002, Czernyadjeva, LE; Sakhalin Region, 23 VIII 2006, Nyushko, MHA.
Ulota hutchinsiae (Smith) Hammar, 1852, Monogr. Orthotrich. Ulot. Suec.: 27. (Fig. 5) Representative illustration: Ignatov et al, 2018 (in prep.): Fig. 63.
Differentiation. Due to saxicolous growth and straight leaves, this species can be easily delineated from the other Ulota species known to occur in Russia, while superficial stomata differ it from species of the genus Orthotrichum, including saxicolous O. anomalum
Fig. 5. Distribution of Ulota hutchinsiae (hollow circle) and U. japonica (solid circle) in Russia.
Hedw. and O. urnigerum Myrin, which prefer calcareous rocks, while U. hutchinsiae mostly occurs on acidic ones. Most similar species of the genus Lewinskya F. Lara, Garilleti et Gof-finet are either predominantly corticolous or have smooth to weakly furrowed capsules and reduced endostome. Anyway, in any case of hesitation, cell pattern in basal leaf portion is useful to segregate these species from Ulota. From the predominantly saxicolous U. curvifolia, U. hutchinsiae differs in having nearly straight leaves and exostome papillose throughout.
Distribution and ecology. Ulota hutchinsiae shows a mostly amphiatlantic suboce-anic pattern of distribution, though it penetrates in Pacific region as well. It is widespread in Britain and southern Scandinavia, while in other European countries it is rarer, gradually declining eastward. In eastern Europe it occurs in Carpathians, Baltic countries, Caucasus and Transcaucasian region. In North America it occurs mostly along Atlantic coast from northern Quebec to Georgia and Alabama; it penetrates westward to eastern Ontario with isolated localities in Arizona and South Alaska (Vitt, 2014). According to Suzuki and Iwatsuki (2013), U. hutchinsiae was recently found in Japan. In western part of European Russia (islands and coastal area of Baltic Sea, southern part of the Republic of Karelia and western Caucasus) it reaches eastern extremity of its Eurasian distribution and occurs in the areas with mild climatic conditions and suitable substrate. It is also documented from a highly disjunctive locality in Altai Mts., middle course of Biya River (Bardunov, 1974; Ignatov, Ochyra, 1994), and since this kind of disjunction turns to be not unique in Ulota, we do not hesitate in the correctness of the identification, despite Wang and Jia (2012) did not include this species in their taxonomic account of the Asiatic Ulota species.
Saxicolous on rather dry acidic crystalline rocks, mostly on gneiss and granite at lower altitudinal belt.
Selected representative specimens: Russia, Leningrad Region, 22 VI 2002, Kurbatova, LE; Krasnodar Territory, 29 VII 2008, Akatova, MHA; Republic of North Ossetia-Alania, 11 X 2013, Khetagurov, LE.
Ulota intermedia Schimp., 1876, Syn Musc. Eur. (ed. 2): 305. (Fig. 6)
Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 64.
Differentiation. This species differs from the other Ulota species known in Russia due to a combination of wide, rounded to obovate, notably concave leaf base, abruptly narrowed to linear-lanceolate acumen, strongly crispate leaves in dry condition, and funnel-shaped or short-cylindrical capsules, not contracted below the mouth. Among other distinctive characters of the species, wide border of short rectangular cells in basal leaf portion, rather narrow yellowish exothecial bands composed of cells with incrassate transverse walls but hyaline lumens, and endostome composed of cells with thickened transverse walls should be listed (cf. Caparros et al., 2016). In strongly crispate leaves and capsules not contracted below the mouth U. intermedia resembles one more species of U. crispa s. l. affinity — U. obtusiuscula Müll. Hal. et Kindb. This species is known to be an endemic of the Pacific North America and is nicely addressed by Persson (1947). According to
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Fig. 6. Distribution of Ulota intermedia in Russia.
Persson (1947) and Vitt (2014), it differs from U. crispa (in fact it rather resembles U. intermedia) in larger size (stem up to 4 cm vs. up to 2.5 cm; setae mostly ca. 4 mm or longer vs. shorter than 3 mm). Few Russian specimens of U. intermedia from Botchi State Reserve (Khabarovsk Territory, Sea of Okhotsk coastal area) comprise rather large plants with stems up to 3 cm high and setae 3-4 mm, but still remarkably smaller than U. obtusi-uscula (two specimens from British Columbia were studied in LE). In our opinion, the status of U. obtusiuscula needs further study. Based on morphology, it can be considered as a variety of U. intermedia despite the phylogenetic tree topology published by Caparros et al. (2016), since only one specimen of U. obtusiuscula was used in this study; U. intermedia clade in that study splits on the next node after U. obtusiuscula and statistical support of the former node is not high. Apparently, due to the obtained topology Caparros et al. (2016) did not considered U. obtusiuscula to be conspecific with U. intermedia, despite the stem, leaf and setae measurements provided in Vitt (2014) for U. obtusiuscula and Caparros et al. (2016) for U. intermedia mostly overlap. On the other hand, the studied U. obtusiuscula specimens share not characteristic to U. intermedia red coloration of the uppermost capsule part, which thus could be considered as the diagnostic feature of the species.
Distribution and ecology. Ulota intermedia has rather wide amphioceanic cool temperate/hemiboreal, moderately oceanic distribution which is still underestimated since for a long time it has been considered as conspecific with U. crispa. According to Ca-parrós et al. (2016), it is widespread along Atlantic and Pacific coasts within Holarctic and mostly does not penetrate far in inland areas of Eurasia excepting the isolated (relic?) population in Altai Mts. According to our results, this is one of the most widespread Ulota
species in Russia. All specimens of Ulota crispa complex in Middle European Russia (cf. Ignatov, Ignatova, 2003; Ignatov et al., 2006; Ivanov et al., 2017) in fact represent U. intermedia; eastward it reaches the Ryazan, Vladimir and Kostroma regions. In north Caucasus it occurs along Main Caucasian Range and, unlike other Ulota species, penetrates also in rather dry foothill area (western extremity of the Kalmykia Republic). It is also widespread in Altai Mts., occurring in places with rather mild and wet local climatic conditions around Teletskoe Lake (Ignatov, Ochyra, 1994), while the specimens from Kulumys Range (Western Sayan Mts.), which were previously identified as U. crispa (Bardunov, 1974) we referred to U. rehmannii, not to U. intermedia. Most specimens previously referred to U. crispa from the coastal areas of the Russian Far East also represent U. intermedia. It sporadically occurs in western, middle and southern parts of the Kamchatka Peninsula and is widespread along Sea of Okhotsk coast within the Khabarovsk and Primorye territories, declining westward and reaching Khabarovsk City area and Badzhal Mts. (ca. 200 km northward from Khabarovsk City), in Sakhalin Island and South Kurils (Fig. 7). Northern limit of the species distribution in the area of Sea of Okhotsk is still unknown due to extremely low degree of bryofloristic exploration of its continental coast. Corticolous, on Populus, Salix, Betula, Picea, Abies, Acer, Pinus pumila, etc.
Selected representative specimens: Russia, Leningrad Region, 27 VII 2011, Kushnevskaya, LE; Novgorod Region, 15 VI 2001, Andreeva, LE; Pskov Region, 6 XI 1996, Ukminskaya, LE; Tver Region, 3 VIII 1994, Ignatov, MW; Moscow Region, 19 VI 2005, Ignatova, MW; Vladimir Region, 8 IX 2008, Kokoshnikova, MW; Ryazan Region, 4 V 2000, Volosnova, MW; Krasnodar Territory, 24 VI 2007, Kurbatova, LE; Republic of Kalmykia, 19 V 2010, Ukrainskaya, LE; Republic of Altai, 24 VI 1991, Ignatov, MHA; Kamchatka Territory, 22 VII 2004, Neshataeva, LE; Khabarovsk Territory, 23 VIII 2013, Ignatov, Ignatova, MHA; Primorye Territory, 15 IX 1947, Rosenberg, LE; Sakhalin Region, 10 IX 2006, Ignatov, MHA.
Ulota japonica (Sull. et Lesq.) Mitt., 1891, Trans. Linn. Soc. London, Bot. 3: 162. (Fig. 5) Representative illustration: Ignatov et al, 2018 (in prep.): Fig. 65.
Differentiation. According to Wang and Jia (2012), the most prominent characters of Ulota japonica are rather small plant size and rectangular, rather thin-walled basal leaf cells near costa with neither sinuose nor porose longitudinal walls. U. barclayi (see above) shares these characters, but differs in obtuse to rounded (vs. acute) perichaetial leaves, oblong to funnel-shaped, brown (vs. mostly urceolate, light brown) capsules and not fused (vs. fused) exostome teeth. According to Wang and Jia (2014), in some populations of U. japonica the cell pattern in basal leaf portion is weaker expressed, and few Russian specimens correspond to this notification; in such a case, it can be confused with U. reptans due to combination of rather small plant size, slightly curved leaves, papillose peristome and sparsely hairy calyptra, though the basal leaf cells in any case are not similar to those in U. reptans. Among other distinctive characters, larger leaves (1.5-2.5 mm vs. 1.0-1.5 mm long) and smaller spores (15-24 pm vs. 27-40 pm) suggest toward U. japonica.
Distribution and ecology. The distribution of U. japonica is amphipacific, rather strictly oceanic, temperate/hemiboreal, and our data contribute to this pattern. Previously the species has been known from Korea, Japan, South Kurils, Sakhalin Island, South Alaska and British Columbia (Noguchi, Iwatsuki, 1989; Bakalin et al, 2009; Wang, Jia, 2012; Bakalin et al., 2014; Vitt, 2014). Here we add localities in the Primorye and Khabarovsk territories and the Kamchatka Peninsula, thus partly filling the gap in the species range in North Pacific. The species seems to be not rare in coastal areas of the Primorye Territory and in South Kuril Islands, where it occurs in lower elevations, growing on willow, poplar, spruce and fir trunks and branches. In the Kamchatka Peninsula U. japonica was found to be quite abundant in Kronotsky State Reserve, where it grows in coastal area on birch and alder trunks and branches. In Iturup Island it was once collected on coastal volcanic rocks.
Selected representative specimens: Russia, Kamchatka Territory, 24 IX 2012, Fedosov, MW; Khabarovsk Territory, 20 VIII 2013, Ignatov, Ignatova, MHA; Primorye Territory, 15 IX 2013, Ignatov, Ignato-va, Malashkina, MHA; Sakhalin Region, 3 IX 2013, Bakalin, MHA.
Ulota rehmannii Juratzka, 1864, Verh. Zool.-Bot. Ges. Wien 14: 191. (Fig. 7)
Representative illustration: Ignatov et al, 2018 (in prep.): Fig. 66.
Differentiation. Ulota rehmannii is readily recognized from U. intermedia, with which it often grows together, by its straight or weakly crisped leaves. When nearly mature capsules are available, U. rehmannii differs from U. intermedia due to bright red (vs. mostly not differentiated in color) rim of the operculum. Noteworthy that in herbarium specimens this character often lacks. In addition, U. rehmannii has somewhat shorter leaves and smooth or somewhat hairy calyptra with darker upper portion.
In having weakly curved leaves, cylindrical to funnel-shaped capsules, large spores and conic, nearly naked calyptrae U. rehmannii is most similar and apparently closely related to East Asian U. reptans and West North American U. megalospora Venturi. Differentiation from U. reptans is considered under that species. Despite distinction of U. rehmannii and U. megalospora is not discussed in any taxonomic studies, it seems to be quite actual issue. Besides the similarities listed above, these species share growth pattern, leaf size, exostome teeth striolation, red-rimmed opercula and some other features. Until present, these species were considered as allopatric, since U. rehmannii occurs in Eurasia, while U. megalospora is a North American species, but Suzuki and Iwatsuki (2013) reported the latter species from Japan, where a year before the former one was revealed (Wang, Jia, 2012). We consider the shape of leaf tips, location of stomata and spore size as distinguishing characters of U. rehmannii and U. megalospora, basing on the key and description of U. megalospora provided by Vitt (2014). At the same time, U. rehmannii has mostly acute leaf tips, stomata located in proximal portion of the urn and spores up to 36(40) pm in diameter. Even superficial examination of the description and images of Japanese specimen referred to U. megalospora by Suzuki and Iwatsuki (2013) in terms of distinctions considered above, suggests that the specimen apparently represents U. rehmannii, which was revealed in Japan by Wang and Jia (2012), but omitted in Suzuki and Iwatsuki (2013). In our opinion, the ecology and distribution pattern are contrasting between hyperoceanic, temperate North American U. megalospora and rather continental, predominantly hemiboreal Eurasian U. rehmannii. They are essential in understanding these species, though further study of U. megalospora — U. macrospora Baur et Warnst. — U. rehmannii — U. reptans complex is needed.
Distribution and ecology. Ulota rehmannii has cool temperate/hemiboreal distribution, mostly associated with continental, though rather mild climatic conditions. In Europe the species mostly occurs in its eastern part (Tatra and Carpathian Mountains) though even there it is known only from few localities; till 1994 it has been considered as a rare middle European endemic (Ignatov, Ochyra, 1994). At the same time, recent publications and further exploration of moss flora of Russia have revealed that it is much more common in Asia than it was considered before. To start with, Ignatov and Ochyra (1994) found that the species is not rare in locally humid conditions in taiga belt (ca. 400-1000 m) near Te-letskoe Lake in Altai Mts. — the area situated well distant from both Atlantic and Pacific coasts. It was also reported from Kulumys Range, Western Sayan Mts. (Ignatov, Ochyra, 1994). The revision of Wang and Jia (2012) revealed U. rehmannii in Changbai Mt. (Jilin, NE China) and Japan, while Pisarenko (2015) found it in Kuznetskij Alatau and Shoria Mts., south Siberia. Furthermore, in IRK we found the specimen referred to this species from Khamar-Daban Range near Baikal Lake. Our revision added numerous localities of
the species in the Russian Far East. In particular, in the Khabarovsk Territory, Badzhal Mts., in Yarap River valley it is the most common corticolous Ulota species, while the most widespread in Russia U. intermedia was found only twice there on Pinus pumila. In Zeya State Reserve, ca. 300 km westward (Amur Province) U. rehmannii is the only corticolous species of the genus. It is quite abundant there. In our opinion South Siberian and Far Eastern locally humid boreal mountain forests at lower elevation belts represent the most suitable conditions for U. rehmannii.
Selected representative specimens: Russia, Republic of Altai, 2 VIII 1991, Ignatov, MHA; Kemerovo Region, 3 VII 2005, Pisarenko, MW; Krasnoyarsk Territory, 16 VII 2010, Bezgodov, MHA; Khabarovsk Territory, 15 VIII 1997, Ignatov, MHA; Primorye Territory, 3 IX 2006, Ignatov, Ignatova, Cherdantseva, MHA.
Ulota reptans Mitt., 1891, Trans. Linn. Soc. London, Bot. 3: 161. (Fig. 8)
Representative illustration: Ignatov et al., 2018 (in prep.): Fig. 67.
Differentiation. Due to weakly curved leaves, cylindrical to funnel-shaped capsules, large spores and conic, nearly naked calyptrae U. reptans is most similar to U. rehmannii and U. megalospora. In separating it from U. rehmannii we follow Wang and Jia (2012). Despite U. reptans is usually differentiated from U. rehmannii mostly in smaller size (leaves mostly 1.0-1.5 mm vs. 1.5-2.3 mm), red-rimmed capsules and peristome ornamentation (mostly papillose vs. striolate), these distinctions are vague. While examining numerous collections from the Russian Far East, we found that few distinctive characters vary a lot and occur in different combinations, thus it turns to be hard to refer several specimens either to one or to another species. In most of specimens both leaf length and exostome
striolation fit well to U. rehmannii, and we refer them to this species despite non semitransparent apical part of exostome teeth (emphasized by Wang and Jia, 2012) — a character that is hard to interpret. Several specimens fit to U. rehmannii in leaf length, while their exo-stome teeth are weaker striolate, with loose papillae. Since in other respects these specimens are identical with typical U. rehmannii, we refer them to this species. The same case was with smaller plants, which share with U. rehmannii remarkably striolate exostome, but in leaf length fit rather to U. reptans or have intermediate leaf length rates. Thus, we refer to U. reptans only few specimens which fit the description of Wang and Jia (2012) in all respects. Since only a few specimens were examined, we found premature to estimate additional distinctive characters among the species or introduce taxonomical novelties. Thus, we decided to follow Wang and Jia (2012) in accepting U. reptans for a while, though consider it as a critical species needed to be tested employing the molecular-phylogenetic approach. The other closely related species, U. megalospora differs from U. reptans in having larger size, striolate exostome teeth, larger spores and apparently strictly North American distribution (see comments to U. rehmannii).
Distribution and ecology. According to the present data, the distribution of U. reptans is cool temperate/hemiboreal, moderately oceanic. Wang and Jia (2012) considered U. reptans to be Japanese endemic, and since their treatment was published, the present localities are the first outside this insular country. In Japan the species occurs in Hokkaido and Honshu (Noguchi, 1989), while Russian specimens originate from Sakhalin Island and the Khabarovsk Territory (Fig. 8), thus reflecting hemiboreal rather than temperate pattern, which can be caused by hot and dry summer in inland monsoon climate of the Primorye Territory. In general, distribution of U. reptans in the Russian Far East is nearly identical with that of U. rehmannii, thus again challenging its specific status. Most specimens cited in literature (i.e., Bakalin et al., 2009; Fedosov et al., 2016) in fact represent U. rehmannii. Corticolous, in Russia occurs in hemiboreal forests on spruce, fir, alder.
Selected representative specimens: Russia, Khabarovsk Territory, 5 VIII 1997, Iwatsuki, MHA; Sakhalin Region, 21 VIII 2006, Ignatov, Teleganova, MHA.
Biogeographical comments on the Holarctic Ulota species
The genus Ulota is known to include bright icons of oceanic distribution. Considered species demonstrate different patterns of association with oceanic environments; since Russian territory is mostly covered by continental climates and only limited patches under oceanic cool temperate and hemiboreal climates occur, some hyperoceanic species do not penetrate here, but gradient of continentality possesses us to grade the species in their relation to moisture/oceanicity. Without doubts, the strictest oceanic species is U. calvescens Wilson, which declines, not reaching Russian border. Then U. drummondii, U. bruchii, and U. hut-chinsiae penetrate to the Kaliningrad and Leningrad Regions and even the Republic of Karelia (the latter species), U. coarctata, U. crispula, U. crispa and U. hutchinsiae — to Western
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Caucasus, but do not extend to the mainland European Russia, where only U. curvifolia (Republic of Karelia, Kola Peninsula, Urals) and U. intermedia occur, however the latter species declines eastward. In the Caucasus U. intermedia, U. crispa and U. crispula reach the Kabardino-Balkarian Republic, while U. hutchinsiae and U. coarctata decline faster, reaching only Teberda mountain area. In Altai Mts. due to locally high moisture five Ulota species occur (Fig. 9), including U. rehmannii, which does not occur in European Russia, and four species are common in the West Caucasus (excepting U. coarctata and U. crispa). In somewhat drier, though still humid areas of West Sajan only three species, U. intermedia, U. rehmannii and U. curvifolia occur, and only U. rehmannii penetrates to Kuznetskij Alatau, Shoria Mountains (Pisarenko, 2014) and Khamar-Daban Range near Baikal Lake, thus in South Siberia U. rehmannii is the most common species of the genus after U. curvifolia. In the Russian Far East four corticolous Ulota species are common: U. drummondii, U. orientalis, U. pacifica and U. japonica mostly occur along coasts and in insular part, U. intermedia, U. crispa and U. crispula penetrate somewhat farer inland and are somewhat less abundant in coastal areas, and finally U. rehmannii reaches rather continental Tukuringra Range in the Amur Region westward and is especially abundant there, so further localities of the species westward are quite expected.
Thus, the grade of species according to their relation to oceanicity/continentality should be as follow: U. calvescens — U. drummondii, U. coarctata, U. bruchii, U. japonica,
Fig. 9. Representation of the genus Ulota (number of species) in regional floras of Russia. White area (without numbers) indicates regions where no one Ulota species has been found. The question mark indicates areas, where representation of the genus is still unconfirmed, but we expect that some Ulota species occur there.
U. orientalis, U. pacifica — U. hutchinsiae (?) — U. crispa, U. crispula — U. intermedia — U. rehmannii — U. curvifolia. Among these, U. bruchii and U. coarctata have amphiatlantic distribution, U. intermedia, U. crispa, U. crispula, U. drummondii, U. hutchinsiae — amphioceanic one, U. japonica is amphipacific, U. rehmannii demonstrates somewhat uncommon disjunctive Eurasian distribution; U. curvifolia is the only species of the genus in Holarctic with not disjunctive circumpolar distribution. The species which occur only in one center of diversity of the genus (cf. Garilleti et al. 2015) are associated with hy-peroceanic temperate climates. They are U. calvescens in Europe, U. obtusiuscula and U. megalospora in Western North America (for comments on finding of this species in Japan see discussion under U. rehmannii). They do not penetrate to Russia, though hy-peroceanic Plenogemma phyllantha (Brid.) Plásek, Sawicki et Ochyra with its dioicous sexual condition, extremely rare sporophyte production and dispersal mostly by gemmae does. This can be explained by its ability to inhabit rocks and thus penetrate along treeless Aleutians westward. One more reason of such limited distribution pattern in some species can be in association with warmer climates, which makes the areas suitable to colonize much more distant, or in difference between temperate climates of eastern and western continental coasts.
Remarkably, despite cool temperate/hemiboreal oceanic distribution patterns, well expressed in most of the Holarctic Ulota species, no one demonstrates remarkable West-Western or East-Eastern disjunction, considered by Schofield (1988) and quite widespread among the bryophytes of oceanic climates. Interestingly, in closely related genera of Orthotrichaceae both West-Western (Orthotrichum pulchellum Brunton, O. columbicum Mitt., see Medina et al., 2012, Pulvigera lyellii (Hook. et Taylor) Plásek, Sawicki et Ochyra, Plenogemma phyllantha, Lewinskya affinis (Schrad. ex Brid.) F. Lara, Garilleti et Goffnet) and East-Eastern (Orthotrichum ohioense Sull. et Lesq., O. pusillum Mitt.) disjunctions occur. This may indicate rather recent radiation of the genus Ulota, at least in Holarctic. Thus, Holarctic Ulota species often occur in different climatic conditions along the coast of the same ocean, but do not occur in nearly the same climatic conditions along the coast of different oceans.
Our results underlined the cool temperate extension of North East Asian center of diversity of the genus Ulota, where two new species were revealed. Remarkably, both of them represent the group of species related to Ulota japonica, which has never undergone circumstantial revision, thus its diversity apparently is underestimated. Diversification of this group as well as group of species related to U. rehmannii & U. megalospora are especially characteristic to north Pacific centres of diversity of the genus.
At the same time, north Pacific sector is still weakly explored and keeping poorly understood taxa of Ulota. At first, rather widespread species of U. crispa complex discussed under U. crispula, which combines characters of all three, U. crispa, U. crispula and U. intermedia is to be mentioned. Status of U. longifolia and U. reptans is still waiting to be proved; and variation of capsule shape of U. intermedia within its area also seems
abnormal. In particular, one specimen from the Kamchatka Peninsula combines crisped acuminate leaves and fusiform capsules, distinctly narrowed toward the mouth. In some respects it corresponds to the recently described East Asian U. delicata, but cell pattern, acuminate leaf apices, finely papillose exostome teeth and smaller spores reject this affinity. Since the specimen is poor and lacks well preserved peristomes, we would prefer to keep it for a while, despite it obviously represents one more new taxon.
Acknowledgments
The authors are grateful to curators of herbaria MW, MHA, LE, NSK, IRK, UUH, VLA, and VBGI for loan of specimens or for assistance in situ, to Ricardo Garilleti and Francisco Lara for helpful comments on the specimens, manuscript and English improving, and to Sergei Dudov for the help with map preparation. The work was supported by Grant # 14-50-00029 «Scientific basis of the national biobank-depository of the living systems» (branch «Plants») from Russian Science Foundation (RNF).
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Литература
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Noguchi A., Iwatsuki Z. 1989. Illustrated moss flora of Japan Vol. 3. Nichinan: 492-742.
Persson H. 1947. Further notes on Alaskan-Yukon Bryophytes. Bryologist 50(3): 279-310.
Pisarenko O. Yu. 2014. Mosses of Salair-Kuznetsk Region (Altai-Sayan Mountain Country) and adjacent plains of West Siberia. Arctoa 23: 33-62.
Plásek V., Sawicki J., Ochyra R., Szczecinska M., Kulik T. 2015. New taxonomical arrangement of the traditionally conceived genera Orthotrichum and Ulota (Orthotrichaceae, Bryophyta). Acta Musei Silesiae, Scientiae Naturales 64: 169-174.
Plásek V., Smoczyk M., Ochyra R. 2016. Rediscovery of the epiphytic moss Ulota coarctata (Orthotrichaceae) in Poland. Polish Bot. J. 61(1): 99-105.
Ros R. M., Mazimpaka V., Abou-Salama U., Aleffi M., Blockeel T. L., Brugués M., Cros R. M., Dia M. G., Dirkse G. M., Draper I., El-Saadawi W., Erdag A., Ganeva A., Gabriel R., González-Mancebo J. M., Granger C., Herrnstadt I., Hugonnot V., Khalil K., Kürschner H., Losada-Lima A., Luis L., Mifsud S., Privitera M., Puglisi M., Sabovljevic M., Sergio C., Shabbara H. M., Sim-Sim M., Sotiaux A., Tacchi R., Vanderpoorten A., Werner O. 2013. Mosses of the Mediterranean, an annotated checklist. Cryptog. Bryol. 34(2): 99-283.
Schofield W. B. 1988. Bryophyte disjunction in Northern Hemisphere: Europe and North America. Bot. J. Linn. Soc. 98(3): 211-224.
Smith A. J. E. 2004. The moss flora of Britain and Ireland. Cambridge: 1012 p.
Smith A. J. E., Hill M. O. 1975. Taxonomic investigation of Ulota bruchii Hornsch. ex Brid., U. crispa (Hedw.) Brid. and U. crispula Brid. I. European material. J. Bryology 8: 423-433.
Suzuki T., Iwatsuki Z. 2013. Notes on Ulota in Japan. Hattoria 4: 99-106.
Tropicos, 2017: http://www.tropicos.org/namesearch.aspx (Date of access: 20 IX 2017).
Vitt D. H. 2014. Orthotrichaceae. Flora of North America North of Mexico. Vol 28: Bryophyta, part 2. New York; Oxford, 37-82.
Wang Q.-H., Jia Y. 2012. A taxonomic revision of the Asian species of Ulota Mohr (Orthotrichaceae). Bryologist 115(3): 412-443.
