CC BY
66Three new cleistocarpous species of the genera Tortula and Microbryum
(Pottiaceae, Bryophyta)
T. N. Otnyukova
Krasnoyarsk, Russia tnotnuykova@gmail.com
Abstract. Three new cleistocarpous species belonging to the genera Tortula and Microbryum were found among specimens in herbaria LE and MHA identified as Pterygoneurum kozlovii (redefined as Pterygoneurum sibiricum), Tortula acaulon, Microbryum vlassovii and Microbryum curvicollum. The new species Tortula arenaria found in collections from the Republic of Dagestan is similar to M. curvicollum in having curved costa but differs in large and roughly papillose spores (vs. smaller and smooth in M. curvicollum), branched papillae (vs. simple papillae in M. curvicollum). The new species Microbryum lydiae found in collections from the Republic of Armenia is similar to M. vlassovii in having ventral outgrowths of costa presented as elongated parenchymatous cells but differs in long-mam-millose leaf lamina and costa cells (long-mammillose-papillose in M. vlassovii); smaller, warty papillose spores (vs. larger, streaky papillose in M. vlassovii). The new species Tortula irinae is similar to cleistocarpous species of the genus Pterygoneurum in having ventral outgrowths of costa, but differs in outgrowths presented as a pad of large, elongated parenchymatous cells (vs. as low lamellae of several rows of small cells in height in P. kozlovii and high lamellae in P. sibiricum). By ventral outgrowths of costa presented as elongated parenchymatous cells it is also similar to some species of the genus Microbryum, from which it differs in leaf lamina and costa cells with several branched papillae (vs. long-mammillose-papillose leaf lamina and costa cells with one branched papilla in M. vlassovii and long-mammillose leaf lamina and costa cells with one large mammilla in M. lydiae). Tortula irinae resembles T. acaulon in having cleistocarpous immersed capsule. Tortula acaulon is a highly variable species represented by several morphotypes of uncertain taxonomic status but strongly differing in many characters of sporophytes and gametophytes. The most important diagnostic characters are the degree of papillae expression and the shape of leaf and costa papillae. By the degree of papillae expression and the shape of papillae, several morphotypes of T. acaulon were revealed: 1 - morphotype without papillae, 2 - morphotype with simple papillae, 3 - morphotype with branched papillae. Tortula irinae differs from T. acaulon morphotypes in having ventral outgrowths of costa lacking in T. acaulon morphotypes. However, of all T. acaulon morphotypes studied, T. irinae can be most probably confused with T. acaulon "branched papillae morphotype" in having strongly branched papillae of costa and leaf cells but easily differing in shape of ventral costa cells protruding as a pad of large, elongated parenchymatous cells in T. irinae vs. ventral costa cells smaller, not protruding in T. acaulon "branched papillae morphotype". Tortula irinae is widespread in Asian part of Russia [Chukotka Autonomous Area, Republic of Sakha (Yakutia), Trans-Baikal Territory]. It was also found in kept in LE moss collections from Asia (Khazakhtan, Mongolia) and Europe (Ukraine).
Keywords: Microbryum curvicollum, Microbryum lydiae, Microbryum vlassovii, Phascum cuspi-datum, Phascum piliferum, Pterygoneurum kozlovii, Pterygoneurum sibiricum, Tortula acaulon, Tortula arenaria, Tortula irinae, geography, morphology, moss flora, taxonomy, Russia [Chukotka Autonomous Area, Republic of Dagestan; Republic of Karachayevo-Circassian, Republic of Sakha (Yakutia), Stavropol Territory, Trans-Baikal Territory]; Armenia, Austria, Azerbaijan, Germany, Kazakhstan, Mongolia, Poland, Romania, Tajikistan, Ukraine.
https://doi.org/10.31111/nsnr/2020.542.515
515
Три новых клейстокарпных вида родов Tortula и Microbryum (Pottiaceae, Bryophyta)
Т. H. Отнюкова
Красноярск, Россия tnotnuykova@gmail.com
Резюме. Выявлено три новых для науки вида из родов Tortula и Microbryum в гербариях LE и MHA, определенных как Pterygoneurum kozlovii, Tortula acaulon, Microbryum vlassovii и Microbryum curvicollum. Новый вид Tortula arenaria, обнаруженный в коллекциях мхов из Республики Дагестан, сходен с M. curvicollum согнутой ножкой, отличается крупными, грубо-папиллозны-ми спорами (мелкие и гладкие у M. curvicollum), разветвленными папиллами (простые у M. cur-vicollum). Новый вид Microbryum lydiae, обнаруженный в коллекциях мхов из Армении, сходен с M. vlassovii наличием брюшных выростов жилки, образованных удлиненными паренхима-тическими клетками, отличается длинномаммиллозными клетками жилки и пластинки листа (длинномаммиллозно-папиллозные у M. vlassovii); более мелкими, бородавчато-папиллозны-ми спорами (более крупные, штриховато-папиллозные у M. vlassovii). Новый вид Tortula irinae сходен с клейстокарпными видами рода Pterygoneurum брюшными выростами жилки, но отличается тем, что выросты образованы крупными, удлиненными паренхиматическими клетками с разветвленными папиллами (выросты образованы низкими ламеллами из нескольких рядов мелких клеток в высоту у P. kozlovii и высокими ламеллами у P. sibiricum). Брюшными выростами, образованными удлиненными паренхиматическими клетками, вид также сходен с клей-стокарпными видами родов Microbryum и Tortula, от которых отличается клетками пластинки листа и жилки с несколькими разветвленными папиллами (длинномаммиллозно-папиллоз-ные клетки пластинки листа и жилки с одной крупной разветвленной папиллой у M. vlasso-vii и длинномаммиллозные клетки пластинки листа и жилки с одной крупной маммиллой у M. lydiae). Tortula irinae сходен с T. acaulon клейстокарпными, погруженными в перихециаль-ные листья коробочками. Tortula acaulon очень вариабельный вид, представленный несколькими морфотипами неопределенного таксономического статуса, но строго различающимися по многим признакам спорофита и гаметофита. Наиболее значимыми диагностическими признаками являются: степень выраженности папилл и форма папилл. По степени выраженности и форме папилл выделено несколько морфотипов T. acaulon: 1 - морфотип без папилл, 2 -морфотип с простыми папиллами, 3 - морфотип с разветвленными папиллами. Tortula irinae отличается от всех морфотипов T. acaulon брюшными выростами жилки, отсутствующими у T. acaulon. Tortula irinae можно спутать только с морфотипом T. acaulon с разветвленными па-пиллами, от которого этот вид легко отличается формой брюшных клеток жилки, у T. irinae эти клетки крупные и выступающие как выросты, в то время как у морфотипа T. acaulon с разветвленными папиллами брюшные клетки жилки мелкие и не выступающие. Tortula irinae широко распространена в Сибири [Чукотский автономный округ, Республика Саха (Якутия), Забайкальский край]. Вид также обнаружен в хранящихся в LE коллекциях мхов из Азии (Казахстан, Монголия) и Европы (Украина).
Ключевые слова: Microbryum curvicollum, Microbryum lydiae Microbryum vlassovii, Phascum cuspidatum, Phascum piliferum, Pterygoneurum kozlovii, Pterygoneurum sibiricum, Tortula acaulon, Tortula arenaria, Tortula irinae, география, морфология, таксономия, флора мхов, Россия [Забайкальский край, Республика Дагестан, Карачаево-Черкесская Республика, Республика Саха (Якутия), Ставропольский край, Чукотский автономный округ], Австрия, Азербайджан, Армения, Германия, Казахстан, Монголия, Польша, Румыния, Таджикистан, Украина.
In moss collections kept in LE and MHA three new cleistocarpous species identified as Pterygoneurum kozlovii Laz., Tortula acaulon (With.) R. H. Zander, Microbryum vlassovii (Laz.) R. H. Zander and Microbryum curvicollum (Hedw.) R. H. Zander were found. To verify the generic position of the studied species, a 2% solution of KOH (Zander 1989, 1993) was used. The studied specimens of the two species showed a yellow KOH reaction, which may indicate that they can belong to the genus Tortula Hedw. rather than to the genus Microbryum Schimp., the third species had a red reaction belonging to the genus Microbryum.
The study of specimens defined as Pterygoneurum kozlovii (redefined as Pterygoneurum sibiricum Otnyukova) led to the discovery of a new species belonging to the genus Tortula Hedw. In one specimen identified as P. kozlovii (later attributed to P. sibiricum) (Otnyukova, 2020) there was an admixture marked as Phascum cuspidatum Hedw., which in present paper attributed to belonging to a new species of the genus Tortula. So, some specimens kept in LE as Tortula acaulon (Phascum cuspidatum and P. pili-ferum Hedw.) were studied. New Tortula species is characterized by yellow KOH reaction; as well as ventral outgrowths of costa lacking in T. acaulon. It is similar to cleistocarpous species of the genus Pterygoneurum Jur. in having outgrowths, but differs in that outgrowths consist of one row of large elongated parenchymatous cells with several strongly branched papillae (vs. formed as lamellae consisting of several rows of small cells in height in P. sibiricum and P. kozlovii).
Due to the fact that Microbryum vlassovii also has ventral outgrowths of costa, all specimens of M. vlassovii kept in LE were studied and the study led to the discovery of another new species of the genus Microbryum. New Microbryum species is characterized by red KOH reaction; it is similar to M. vlassovii in having ventral outgrowths formed by one row of large elongated parenchymatous cells, but differs in that cells long-mammillose (vs. long-mammillose-papillose in M. vlassovii). It is also characterized by smaller, warty-papillose spores (vs. larger, streaky papillose in M. vlassovii).
In one specimen of Tortula acaulon in LE there was Microbryum admixture, which was marked as "Microbryum curvicolle" (M. curvicollum), but the admixture was redefined as belonging to new species of the genus Tortula. So, all specimens of M. curvi-collum kept in LE were studied. New Tortula species is characterized by bright-yellow KOH reaction; it is similar to M. curvicollum in having costa curved but differs in large, rough papillose spores (vs. smaller, smooth in M. curvicollum). It is also characterized by globose capsules (vs. oval in M. curvicollum); branched papillae (vs. simple in M. curvicollum).
Thus, all cleistocarpous specimens studied are morphologically distinct and deserve to be separate species.
Tortula arenaria Otnyukova, sp. nov. (Plate I)
Diagnosis. Tortula arenaria is similar to Microbryum curvicollum in having strongly curved costa but differs in large and roughly papillose spores (vs. smaller and smooth
Plate I. Tortula arenaria Otnyukova, sp. nov. 1, 3, 4, 6, 8 — capsules; 2, 7 — spores; 5 — calyptra; 9-16 — leaves; 17-21 — leaf cross sections; 22 — upper part of leaf lamina with marginal papillae; 23 — leaf cells with branched papillae; 24, 25 — branched leaf papillae, mainly C-shaped with auricles (in contour). Scale bars: 1, 3-6, 8-16 — 1 mm; 2, 17-21 — 80 |m; 7 — 40 |m; 22, 23 — 25 |m; 24, 25 — 8 |m. From holotype (17 V 2009, G. Ukrainskaya 14100/2, LE).
in M. curvicollum); in globose to ovoid capsule (vs. short-ellipsoid in M. curvicollum); and in papillae branched (vs. simple in M. curvicollum).
Type. Russia, Republic of Dagestan, Sary-Kum dune, on soil, 17 V 2009, M. Ignatov, E. Ignatova, G. Ukrainskaya (М. Игнатов, Е. Игнатова, Г. Украинская) 14100/2 (det. by G. Ukrainskaya as Microbryum curvicolle being in admixture to Tor-tula acaulon), LE (holotype).
Description. Plants up to 2 mm, in loose turfs. Stems mainly forked or simple, crowded leafy. Leaves loosely appressed, erect or slightly curved when dry, erect-spreading when moist, 1.6-1.8(2.2) x 0.35-0.45 mm, oblong-lanceolate to ovate-lanceolate, short to ± long acuminate, keeled; margins ± serrulate due to protruding papillae, plane or narrowly recurved. Costa strong, ± long excurrent, 0.5-0.6(1.0) mm, in cross section with one row of guide cells and ± protruding dorsal stereid band; ventral epidermis large-celled, papillose, dorsal epidermis large-celled only near lamina, papillose, and small-celled on the protruding part of costa, epapillose; ventral cells in cross section elongated, barrel-shaped, (18)20-25 x 15-20(25) jm, dorsal cells in cross section isodiametric, 15-18 jm; papillae of costa 2-4 per cell, branched (in the upper part of costa), simple or lacking (in the lower part of costa). Cells of leaf lamina in the upper part isodiametric, quadrate, quadrate-hexagonal,
Plate II. Microbryum curvicollum (Hedw.) R. H. Zander. 1, 4 — spores; 2, 5, 6 — capsules; 3 — leaf; 7, 8 — leaf cross sections; 9, 10 — simple leaf papillae, rounded (in contour); 11 — lamina cells with papillae; 12 — part of leaf lamina with marginal papillae. Scale bars: 1, 4 — 40 |m; 2, 3, 5, 6 — 1 mm; 7 — 80 |m; 8 — 150 |m; 9, 10 — 8 |m; 11, 12 — 25 |im. 1, 2, 4 — from 19 X 1903, C. Jensen, LE; 3, 5-12 — from 23 V 2010, G. Ukrainskaya, LE.
13-15(17) X 13-15 jim, in the middle short-rectangular, (13)15-20 x 13-15 jm, in the lower part rectangular, 25-40 x 15 |im, papillose on both sides above, epapillose below; papillae of lamina 2-4 per cell, in the upper part strongly branched, C- or C-shaped with auricles, in the middle 2-divided or simple, in the lower part lacking. Sexual condition autoicous. Sporophytes one or two per each perichaetium. Seta 0.6-0.8 mm long, (0.1)0.15(0.2) mm thick, strongly curved. Capsule cleistocarpic, globose to ovoid, (0.7)0.8-1.0 x (0.5)0.6-0.7 mm, with indistinct apex. Calyptra up to 0.4 mm, triangular-conical, smooth. Spores (30)32-35(37) |im, ± roughly papillose. KOH color reaction of leaf lamina bright yellow or somewhat yellow-orange in cells of stem and costa.
Etymology. The species name derives from Latin adjective arenarius — inhabiting on sand, sandy.
Distribution. Tortula arenaria is found in Caucasus, it is known only from type locality in Sary-Kum dune in Dagestan.
Ecology. Tortula arenaria grows in dry conditions on soil.
Differentiation. Tortula arenaria is similar to Microbryum curvicollum (Plate II) in having strongly curved costa but differs in many other characters. Both species differ in response to KOH, which is bright-yellow in Tortula arenaria vs. red in Micro-byum curvicollum; in length and thickness of seta, which are 0.6-0.8 mm long and up to 0.2 mm thick in Tortula arenaria vs. 0.8-0.9 mm long and up to 0.1 mm thick in Microbryum curvicollum. Tortula arenaria also differs from Microbryum curvicollum in capsule larger, globose to ovoid, 0.8-1.0 x 0.6-0.8 mm vs. smaller, short-ellipsoid, 0.7-0.8 x 0.4-0.5 mm; spores larger and roughly papillose, 32-35 |im vs. smaller and smooth, 22-25 |im; cells of ventral epidermis much larger, strongly protruding as ventral outgrowths vs. smaller, not differing from surrounding cells; papillae of lamina and costa large, branched, C-shaped with auricles or C-shaped in contour vs. small, simple, rounded in contour in M. curvicollum.
Selected specimens examined. Microbryum curvicollum — Russia, Republic of Kalmykia, Ikiburul District, Cholun-Khomur village, career, on soil among fragments of shell rock, 23 V 2010, G. Ukrainskaya (Г. Украинская) 136-кл (as Microbryum curvicolle), LE; Denmark, Zealand, wood at Allindelille, north of Ringsted, 19 X 1903, C. Jensen (Bryophyta Danica Exciccata #171 as Phascum curvicolle Hedw.), LE; Poland, Naklo nad-Noteciq. Bewohner der pontishen Hügel ostlich von Nakel, aber nicht alljarlich so schön entwickelt. Gesammlet von V. Torka am 13 Februar 1913 in Gemeinschaft von Mildea bryoides (Dicks.) Warnst., Pterygoneurum subsessile (Brid.) Jur., Silene otitis Sm. (Bryotheca Posnaniensis #61 as Phascum curvicollum Ehrh.), LE; Sweden, Gotland, Visby, 28 IV 1900, K. Johansson (Musci Suecici, a Museo Botanico Upsaliensi Distributi as Phascum curvicolle Hedw.), LE; Turkmenistan, Kopet-Dag Range, Parkhai Stream, 9 X 1985, I. V. Sirotina (И. В. Сиротина) (det. by A. L. Abramova as Phascum curvicolle), LE; Ukraine, Askania-Nova Reserve, steppe, 11 V 1923, V. N. Sukachev (В. Н. Сукачев) (det. by L. I. Savicz as Phascum curvicollum), LE; ibidem, 12 V 1923, V. N. Sukachev (В. Н. Сукачев) (det. by L. I. Savicz as Phascum curvicollum), LE.
Microbryum lydiae Otnyukova, sp. nov. (Plate III)
Diagnosis. Microbryum lydiae is similar to Microbryum vlassovii in having ventral outgrowths of costa as large elongated parenchymatous cells but differs in that cells
22 23 25 26 27
Plate III. Microbryum lydiae Otnyukova, sp. nov. (1-17) and Microbryum vlassovii (Laz.)
R. H. Zander (18-27).
1, 20, 24 — capsules; 2, 3, 23 — caliptrae; 4, 5, 20, 27 — spores: warty papillose (5), streaky papillose
(27); 6, 8, 12-19, 25, 26 — leaf cross sections; 7 — leaf tip; 9 — part of leaf; 10, 11, 21 — leaves. Scale bars: 1, 10, 11, 18, 20, 21, 23 — 1 mm; 2, 3, 7, 9, 23 — 0.3 mm; 4, 20 — 25 |m; 5, 27 — 15 |m; 6, 8, 12 — 150 |im; 13, 15 16, 19, 25, 26 — 80 |m; 14, 18 - 20 |m; 17 — 75 |m. 1, 3, 6, 7, 13-16 — from paratype (7 III 1953, A. G. Abramyan, A. M. Bargesyan, LE); 2, 4, 5, 8-12, 17 — from holotype (3 III 1953, A. G. Abramyan, A. M. Bargesyan); 18-27 — from 10 IV 1963, A. L. Lyubarskaya, LE.
long-mammillose (vs. long-mammillose-papillose cells ending in one branched papilla in M. vlassovii); as well as in smaller, warty papillose spores (vs. larger, streaky papillose in M. vlassovii).
Type. Armenia, Yerevan City, Botanical Garden, 3 III 1958, A. G. Abramyan, A. M. Bargesyan (А. Г. Абрамян, А. М. Баргесян) [det. by A. and I. Abramovy as Lyd-iaea vlassovii (Laz.) Laz.], LE (holotype, LE).
Paratype from the type locality. Armenia, Yerevan City, Botanical Garden, Artemisia erevanica grouping, 7 III 1958, A. G. Abramyan, A. M. Bargesyan (А. Г. Абрамян, А. М. Баргесян) (det. by A. and I. Abramovy as Lydiaea vlassovii), LE.
Description. Plants up to 5 mm, scattered or gregarious. Stems simple, kidney leafy. Leaves gradually becoming larger upward, oval-ovoid, (1)2-3 x (0.4-0.7)1.0-1.3 mm, concave; margins entire, widely recurved above, plane below. Costa ± strong, excurrent, 0.4-0.6 mm, in cross section with one row of guide cells and ± strong dorsal stereid band; ventral epidermal cells large, strongly protruding as ventral outgrowths, in cross section conical, long-mammillose, 35-50 jm long; dorsal epidermal cells ± smooth. Cells of leaf lamina isodiametric, quadrate-hexagonal, in the lower part at margins short-rectangular, (13-15)17-20 x 13-17(20) jm, all long-mammillose on ventral side, smooth or somewhat short-mammillose on dorsal side, in the upper part with one large mammilla, in the lower part at margins with (1)2-4 low mammillae (or papillae); near costa somewhat long-rectangular, colorless, 25-40 x 15 jm, smooth. Sexual condition autoicous. Sporophytes immersed. Seta short, 0.2-0.3 mm. Capsule cleistocarpic, short-ellipsoid, 0.6-0.7 x 0.4-0.5 mm, with short, narrowly-triangular blunt apex. Ca-lyptra up to 0.3 mm, conical, smooth. Spores 18-22(25) jm, warty papillose. KOH color reaction of leaf lamina red or somewhat brownish in cells of stem and costa.
Etymology. The species name is given in honor of Lydia I. Savicz-Ljubitskaja (Savich-Ljubitskaya), the famous Russian bryologist (St. Petersburg, Russia).
Distribution. Microbryumlydiae is found in Caucasus, it is known only from type locality in Yerevan Botanical Garden in Armenia.
Ecology. Microbryum lydiae grows in dry conditions on soil among grasses.
Differentiation. Microbryum lydiae is similar to M. vlassovii in having ventral outgrowths of costa presented as elongated parenchymatous cells. Both species differ in shape of cells, which are long-mammillose in M. lydiae vs. long-mammillose-papil-lose in M. vlassovii. Microbryum lydiae also differs from M. vlassovii in capsule small, short-ellipsoid, 0.6-0.7 x 0.4-0.5 mm vs. larger, ellipsoid, 0.9-1.0 x 0.6-0.7 mm; in spores small, warty papillose, 18-22 jm vs. larger, streaky papillose, 25-28 jm; in long-mammillose cells with one large mammilla vs. long-mammillose-papillose cells with one branched papilla in M. vlassovii.
Selected specimens examined. Microbryum vlassovii — Azerbaijan, Masally District, in surroundings of Masally Town, open sites, on soil, 10 IV 1963, A. L. Lyubarskaya (А. Л. Любарская) (det. by A. L. Abramova as Lydiaea vlassovii), LE; Kyrgyzstan, Kyrgyz Ala-Too (Kyrkyzsky Ridge), natural boundary Chon-aryk, north exposure slope, 950 m a. s. l., on soil, 10 V 1965, E. Rakhmatuli-na (Э. Рахматулина) (as Lydiaea vlassovii; in admixture to Pterygoneurum ovatum (Hedw.) Dixon
and Syntrichia sp.), LE; Tajikistan, Dushanbe City, Botanical Garden, on moist soil, 23 III 1963, U. K. Mamatkulov (У. К. Маматкулов) 3985 (as Lydiaea vlassovii), LE; Turkmenistan (Trans-Caspian Region), Mahtumkuli etrap (Karakalinskoye pristavsto), Kopet-Dag Range (Salchy-dag), 22 III 1916, E. G. Tschernjakovskaja (Е. Г. Черняковская) 97 (det. by A. and I. Abramovy in 1963 as Lydiaea vlassovii; in admixture to Phascum acaulon), LE; Uzbekistan, Tashkent Region, Chirchiq River Basin, adyrs, on soil, 17 IV 1963, E. Rakhmatulina (Э. Рахматулина) (det. by A. and I. Abramovy as Lydiaea vlassovii; in admixture to Tortula sp.), LE; Tashkent City, on ground roofs and soil in open spaces, 15 IV 1926, I. Raykov (И. Райков) (det. by A. and I. Abramovy in October 1963 as Lydiaea vlassovii; in admixture with Pterygoneurum ovatum), LE.
Tortula irinae Otnyukova, sp. nov. (Plates IV, V, VI, VII)
Diagnosis. Tortula irinae resembles T. acaulon in immersed cleistocarpous capsule, but differs in having ventral outgrowths of costa lacking in T. acaulon. In ventral outgrowths of costa it is similar to cleistocarpous species of the genera Pterygoneurum, Microbryum and Tortula but differs in having one row of elongated ventral cells (vs. several rows of small cells in height as lamellae in Pterygoneurum sibiricum and P. koz-lovii); from Microbryum and Tortula species with one row of elongated ventral cells it differs in having cells large, barrel-shaped with several branched papillae (vs. cells long-mammillose-papillose in M. vlassovii and long-mammillose in M. lydiae).
Type. Russia, Trans-Baikal Territory, Sokhondinsky State Natural Biosphere Reserve, surroundings of cordon on river Enda, 49°27'N, 110°51'E, 1070 m a. s. l., stony steppe slope, on bare soil among stones, 14 VII 2010, I. V. Czernyadjeva 19-10 (as Tortula acaulon, LE) (holotype, LE).
Paratypes from the same locality. Russia, Trans-Baikal Territory, Sokhondinsky State Natural Biosphere Reserve, surroundings of cordon on river Enda, 49°26'N, 110°48'E, 1473 m a. s. l., steppe slope, on bare soil under stones, 28 VIII 2011, I. V. Czernyadjeva 55-11 (as Tortula acaulon), LE; ibidem 49°26'N, 110°53'E, 1239 m a. s. l., rocky steppe slope, on bare soil, 29 VIII 2011, I. V. Czernyadjeva 57-11 (as Tortula acaulon), LE.
Paratypes from the other territories. Russia, Chukotka Autonomous Area, Central Chukotka, middle course of Palyavaam River, cryophyte steppe with herb and Carex, 21 VII 1989, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum, with capsules; with Bryum argen-teum Hedw.), LE; ibidem, steppe cryophyte grouping on top of outlier, 10 VII 1989, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum; with Bryum argenteum), LE; ibidem, herb-grass cryophyte steppe, 19 VII 1989, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum; with Desmat-odon latifolius (Hedw) Brid. and Stegoniapilifera (Brid.) H.A. Crum & L.E. Anderson), LE; ibidem, 19 VII 1989, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum), LE; ibidem, steppe birch forest on southern slope of hill, 19 VII 1989, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum (marked by O. Afonina as having "low outgrowths on ventral side of costa"); with Bryum argenteum), LE; Republic of Sakha (Yakutia), right bank of Indigirka River, near mouth of Ystan-Yuryakh (opposite village of Tyubelyakh), grass meadow steppe with Helictotrichon and Poa, 1 VIII 1976, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum var. piliferum, with capsules), LE; ibidem, larch forest edge, herb-grass steppe with Lycopodium and epigeous lichens, 29 VI 1976, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum var. piliferum, with capsules; with Bryum argenteum), LE; ibidem, herb-grass mountain steppe with Alyssum, Artemisia, Helictotrichon and Koeleria,
Plate IV. Tortula irinae Otnyukova, sp. nov. 1, 5 — capsules; 2, 9 — calyptrae; 3 — beak of capsule; 4, 10, 16, 20, 21 — stem leaves; 6, 15 — spores;
7 — outer perichaetial leaf; 8 — outer perichaetial leaf with antheridia; 11— antheridia; 12-14, 17-19, 23 — leaf cross sections; 22 — leaf papillae (inprofile); 24 — young leaf with ventral
outgrowths; 25 — part of leaf with ventral outgrowths; 26 — upper part of costa with ventral outgrowths as a pad of elongated cells. Scale bars: 1, 4, 7, 8, 10, 16, 20, 21 — 1 mm; 2, 5 — 0.5 mm; 3 — 300 |m; 6, 15, 17, 26 - 40 |m; 9 - 350|m; 11 — 250 |m; 12-14, 18, 19, 23 — 80 |m; 22 — 20 |m; 24 — 150 |m; 25 — 60 |m. 1-3, 5, 6, 15, 18, 19, 22 — from holotype (14 VII 2010, Czernyadjeva 19-10, LE); 4, 7, 8, 10, 11, 16, 20, 21 — from paratype (21 VII 1989, Afonina, LE); 12-14, 17, 23-26 — from paratype (22 VII 2010, Czernyadjeva37-10, LE).
Plate V. Tortula irinae Otnyukova, sp. nov. 1 — upper part of leaf with ventral outgrowths; 2 — ventral outgrowths in the upper part of leaf; 3-7 — branched leaf papillae, C-shaped and C-shaped with auricles; 8 — leaf lamina cells; 9 — ventral outgrowths in the lower part of leaf. Scale bars: 1, 2, 8, 9 — 25 |im; 3-7 — 8 |im. 1, 2, 4, 5, 8, 9 — from paratype (22 2VII 2010, Czernyadjeva 37-10, LE); 3, 7 — from paratype (21 VII 1989, Afonina, LE); 6 — from holotype (14 VII 2010, I. V. Czernyadjeva 19-10, LE).
Plate VI. Tortula irinae Otnyukova, sp. nov. 1 — leaf cells in the upper part of leaf; 2, 3, 9, 12, 13 — leaf cross sections; 4 — leaf cells in the middle part of leaf; 5 — leaf cells in the lower part of leaf; 6, 7, 11, 15 — branched leaf papillae, C-shaped with auricles (6, 15) and 2-divided (7, 11); 8, 10 — cross section of ventral outgrowths with elongated, barrel-shaped cells; 14 — cells of leaf lamina. Scale bars: 1, 4, 5 — 20 |m; 2, 3, 9, 13 — 80 |m; 6, 7, 11, 15 — 8 |m; 8, 10, 14 — 25 |m; 12 — 150 |m. From paratype (27 VII 1976, M. Boiko, LE).
Plate VII. Tortula irinae Otnyukova, sp. nov. 1 — branched leaf papillae, C-shaped with auricles; 2, 5 — cells of the ventral outgrowths in the upper part of the leaf; 3 — leaf cells in the upper part of leaf; 4, 9 — leaf lamina cross sections; 6 — cells of the ventral outgrowths in the lower part of the leaf; 7 — simple papillae in the low part of leaf; 8 — branched papillae in the middle part of leaf; 10, 11 — costa cross sections. Scale bars: 1, 4, 7, 8 — 8 |m; 2, 3 — 25 |m; 4 — 20 |m; 5, 6 — 40 |m; 9-11 — 80 |m. From paratype (20 III 1877, A. Regel, LE).
on slope of south exposure, 29 VI 1976, O. M. Afonina (O. M. Афонина) [as Phascum cuspidatum var. piliferum (Schreb. ex Hedw.) Hook. et Taylor, with capsules], LE; left bank of Indigirka River, above mouth of Inyali River, 18 VI 1976, O. M. Afonina (O. M. Афонина) (as Phascum cuspidatum var. piliferum, with capsules; with admixture marked by O. Afonina as Pterygoneurum kozlovii (redefined as Pterygoneurum sibiricum) and Bryum argenteum), LE; Oimyakon District, surroundings of Ust-Nera Settlement, 64°32'27.8"N, 142°58'10"E, 585 m a. s. l., left bank of Indigirka River, 957 km along Magadan Road, dry steppe slope, on bare soil among grasses, 31 VII 2015, Ignatov, Ignatova (Игнатов, Игнатова) 15-1020, MHA 9046875; ibidem, 60 km upstream Nera Riverm from Ust-Nera Selllement, 64°27'20"N, 144°21'30"E, 650 m a. s. l., 60 km to east from Ust-Nera Settlement, right bank of Nera River Valley near mouth of Topol' Stream, steppe slope, on soil under stone, 5 VIII 2015, Ignatov, Ignatova (Игнатов, Игнатова) 15-1483, MHA 9046873; Trans-Baikal Territory, Kyrynsky Dictrict, Sokhondinsky State Natural Biosphere Reserve, surroundings of cordon on Agutsa River, 49°27'N, 110°51'E, 1070 m a. s. l., steppe slope, on bare soil, 22 VII 2010, Czernyadjeva (Чернядьева) 37-10 (as Tortula acaulon), LE; ibidem, Enda River, 49°26'N, 110°52'E, 1117 m a. s. l., forb-grass steppe on stony slope, 14 VII 2010, O. M. Afonina (O. M. Афонина) A2510 (as Tortula acaulon), LE; ibidem, Khukhe-Byatsa Creek, 49°27'50"N, 110°52'11" E, 1214 m a. s. l., dry rocky outcrops on S-facing slope, 28 VIII 2011, O. M. Afonina (O. M. Афонина) 7611 (as Tortula acaulon), LE; Ononsky District, near Nizhni Zasuchey, slope of Onon River terrace, 59°32'N, 114°59'E, 737 m a. s. l., 18 VII 2006, O. M. Afonina (O. M. Афонина) 5406 (as Tortula acaulon; with Bryum argenteum), LE; Borsinsky District, Kululbey Ridge, surroundings of Perednaya Byrka village, 50°45'39"N, 116°46'23"E, 880 m s. a. l., rocky outcrops on S-facing slope, on bare soil among stones, 27 VII 2012, Czernyadjeva (Чернядьева) 34-12 (as Tortula acaulon; with Weissia Hedw.), LE; Olovyanensky District, surroundings of Olovyanyi village, 50°59'01"N, 115°35'43"E, 566 m a. s. l., Onon River bank, rocky outcrops on bank, 29 VII 2012, Czernyadjeva (Чернядьева) 36-12 (as Tortula acaulon), LE; Nerchinsko-Kuenginsky Ridge, Uldurga River, a tributary of Nercha River, 52°37'40"N, 115°24'11"E, 620 m a. s. l., rocky outcrops on S-facing slope, in crevices of stones, 17 VII 2012, Czernyadjeva (Чернядьева) 13-12 (as Tortula acaulon), LE; Daursky Ridge, Aratsa River, tributary of Ingoda River, 51°54'50"N, 114°25'06"E, 900 m a. s.l., roadside, on bare soil, 14 VII 2012, Czernyadjeva (Чернядьева) 4-12/2 (indicated by I. Czernyadjeva as Phascum cuspidatum; in admixture with Pterygoneurum kozlovii, redefined as Pterygoneurum sibiricum), LE; Borschovka Ridge, surroundings of Baley Town, near Sukhoi Fall, on stony (granites) outcrops, 12 VII 1957, I. Abramov (И. Абрамов) (as Phascum piliferum), LE; — Khazakhstan, Almaty (Werny), Almatinka minor pr. Werny, 20 III 1877, A. Regel (det. by A. Abramova in 1961 as Phascum cuspidatum), LE; Mongolia, Dorkot ai-mak, Bayan-Dun somon, Sogoot-Ula Mountains, 1220 m a. s. l., on soil, 22 VII 1974, N. Golubkova, U. Tsogt (Н. Голубкова, У. Цогт) (det. by A. and I. Abromovy as Phascum cuspidatum), LE; Ulaan-baatar, Bogdo Ula Mountain, fall Zaisan, NW exposure slope, herb steppe with Festuca, on soil, 12 VI 1974, Ts. Tsegmed (Ц. Цэгмэд) 896 (as Phascum cuspidatum), LE; Ukraine, Kherson Region, Aska-nia-Nova Reserve, steppe, 27 VII 1976, M. Boiko (М. Бойко) (as Phascum piliferum), LE.
Description. Plants up to 2 mm, in loose turfs, scattered or gregarious. Stems crowded leafy. Leaves loosely appressed, curved or slightly twisted when dry, erect-spreading when moist, (0.6-0.9)1.2-1.8(2.2) x 0.4-0.5(0.6) mm, ovate to oblong-lanceolate, ± concave; margins narrowly recurved from base to apex, in the upper part plane, serrulate due to protruding papillae. Costa strong, ± long excurrent, awn 0.4-0.6(1.0) mm; ventral epidermal cells large, strongly protruding as ventral outgrowths, in cross section elongated, barrel-shaped, (25)30-35 x (15)20-25 |im, papillose; dorsal epidermis small-celled, papillose; papillae of costa 2-4 per cell, in the upper part of costa branched, 2-4(5)-divided, in the lower part of cos-
ta simple or lacking. Cells in the upper part of leaf lamina isodiametric, quadrate, 13-15(17) x 13-17 |m, in the middle short-rectangular, 17-20(25) x 13-15 |m, papillose; papillae 1-3(4) per cell, on both sides; in upper lamina cells strongly branched, 2-4(5)-divided, in the middle 2-divided or simple, lower lacking. Sexual condition sinoicous. Sporophytes immersed. Seta very short, 0.05-0.1 mm, at base strongly bulbiform. Capsule cleistocarpic, globose, 0.5-0.6 mm, with distinct short, wide-triangular blunt apex; peristome absent; operculum not differentiated. Ca-lyptra small, 0.2-0.3 mm, narrowly conical, smooth. Spores 20-25 |m, ± densely papillose. KOH color reaction of leaf lamina yellow.
Etymology. The species name is given in honor of Irina V. Czernyadjeva, the bryo-logist from Komarov Botanical Institute, St. Petersburg, Russia, a collector of this species.
Distribution. Tortula irinae is widespread in the Asian part of Russia (Siberia), including Trans-Baikal Territory, Republic Sakha (Yakutia) and Chukotka Autonomous Area. The species also found in Asia (Khazakhtan, Mongolia,) and Europe (Ukraine). It is logical to assume that species is more widespread in the world.
Ecology. Tortula irinae grows on soil in dry conditions. In Siberia it grows on the steppe slopes among grasses. It is very often found together with Pterygoneurum sibiri-cum in separate small turfs or in admixture to mosses.
Differentiation. In cleistocarpous capsules T. irinae resembles T. acaulon, but differs in having ventral outgrowths (Plates IV-VII) lacking in T. acaulon (e. g., Zander, Eckel, 2007), as well as strongly papillose distal laminal cells (Plates IV-VII), weakly simply papillose or smooth (Savicz-Ljubitskaja, Smirnova, 1970 as Phascum cuspida-tum; Ignatov et al, 2003 as T. acaulon; Zander, Eckel, 2007 as T. acaulon).
Tortula irinae may be confused with T. acaulon, a highly variable species represented by several morphotypes of uncertain taxonomic status (Plates VIII-X). These morphotypes were revealed in the course of comparative study of T. irinae in LE collections, briefly characterized and keyed out below. They described here to delineate better the differentiation of T. irinae and deserve further special study.
All cleistocarpous specimens "looking like T. acaulon" studied in LE are strongly distinct in the following contrast characters: ventral outgrowths of costa (present or lacking); the degree of leaf papillae expression (present or lacking); shape of papillae (branched or simple); shape of capsule (spherical or ellipsoid). Length of costa is a variable character. Among specimens "looking like T. acaulon" studied in LE several morphotypes were found, differing in the severity of ventral outgrowths of costa and leaf papillae.
1. Ventral outgrowths of costa present...................................................................T. irinae (Plates IV-VII)
— Ventral outgrowths of costa lacking..............................................................T. acaulon morphotypes (2)
2. Leaf and costa cells smooth...........................................T. acaulon "smooth morphotype" (Plate VIII)
— Leaf lamina and costa cells ± papillose; papillae 1-4 per cell................................................................(3)
3. Papillae simple, sometimes 2-divided..............................................................T. acaulon "simple papillae
morphotype" (Plate IX)
Plate VIII. Tortula acaulon morphotype without papillae ("smooth morphotype"). 1, 2, 18 — capsules; 3-5, 12, 13, 19, 21 — steam leaves; 6, 10, 20, 22 — leaf cells; 7, 11, 14, 16 — spores; 8, 9, 15, 17 — leaf cross sections. Scale bars: 1-5, 12, 13, 18, 19, 21 — 1 mm; 6, 10, 20, 22 — 20 |im; 7, 11, 14, 16 — 40 |m; 8, 9 — 80 |m; 15, 17 — 150 |m. 1-7, 10, 11, 12-14 — from 11 III 1993, G. Ukrainskaya, LE; 8, 9, 15, 17 — from 20 IV 1873, E. Zickendrath, LE; 18-22 — from 14 VIII 1993, G. Ukrainskaya, LE.
— Papillae branched, 2-4(5)-divided.............................................................T. acaulon "branched papillae
morphotype" (Plate X)
Tortula acaulon "smooth-morphotype" (morphotype without papillae) is characterized by lacking ventral outgrowths; leaf and costa cells smooth (Plate VIII).
Tortula acaulon "simple papillae morphotype" (morphotype with simple papillae) is characterized by lacking ventral outgrowths; leaf and costa cells papillose; papillae 1-4 per cell, simple or 2-divided (Plate IX).
Tortula acaulon "branched papillae morphotype" (morphotype with branched papillae) is characterized by lacking ventral outgrowths; leaf and costa cells strongly papillose; papillae 1-4 per cell, branched, 2-4(5)-divided (Plate X).
Of all T. acaulon morphotypes studied, T. irinae can be most probably confused with T. acaulon "branched papillae morphotype" in having strongly branched papillae of lamina and costa cells, but differs in shape of ventral costa cells protruding as a pad of large, elongated parenchymatous cells in T. irinae vs. ventral costa cells smaller, not protruding in T. acaulon "branched papillae morphotype".
Selected specimens of Tortula acaulon examined: morphotype without papillae — Russia, Stavropol Territory, District of Caucausian Mineral Waters, surroundings of city of Kislovodsk, on ground in cracks between stones, 11 VIII 1993, G. Ukrainskaya (Г. Украинская) кмв-32 (as Tortula acaulon), LE 13475; ibidem, surroundings of Gorny village, 43°57' N, 42°51' E, on soil among grounds, 2 V 2009, G. Ukrainskaya (Г. Украинская) (as Tortula acaulon), LE 13939; Karachayevo-Circassian Republic, Bechisyn Plateau, on soil, 14 VIII 1993, G. Ukrainskaya (Г. Украинская) (as Tortula acaulon), LE 15698; — Austria, Vienna (Wien), Mödling (Kreuzberg bei Moedling), 20 IV 1873, Ernst Zickendrath 47 (as Phascum cuspidatum var. piliferum), LE; morphotype with simple papillae — Russia, Stavropol Territory, District of Caucausian Mineral Waters, Beshtau Mountain, Kozii Skaly, on ground in grass, 7 V 2009, G. Ukrainskaya (Г. Украинская) (as Tortula acaulon; with Barbula unguiculata), LE 15287; Khazakhastan, Ak-mola Region (Tselinograd Region), Kogalzhyn village, island in northeast of Tengiz Lake, on soil among Halocnemum, 15 VII 1988, T. V. Sidorova (Т. В. Сидорова) (det. by O. M. Afonina as Phascum cuspidatum var. piliferum), LE; ibidem, Tengiz Lake, northeast coast, on soil among Halocnemum, 15 VII 1984, T. V. Sidorova (Т. В. Сидорова) (det. by O. M. Afonina as Phascum cuspidatum var. piliferum; with Physcomitrium pyriforme), LE; ibidem, 40 km south-southwest of Kogalzhyn village, northeast coast of Kogalzhyn Lake, on ground in meadow community, 30 VI 1979, T. V. Sidorova (Т. В. Сидорова) (as Phascum cuspidatum), LE; Poland (Polonia), Wyzyna Lubelska, Lublin, ad terram loessaceam in oppido, 4 IV 1966, K. Karczmarz (Musci Exciccati Palatinatus Lublinensis-Polonia # 132 as Phascum piliferum Schreb.), LE; morphotype with branched papillae — Russia, Republic of Dagestan, Sary-Kum dune, on soil, 17 V 2009, M. Ignatov, E. Ignatova, G. Ukrainskaya (М. Игнатов, Е. Игнатова, Г. Украинская) 14100 (det. by G. Ukrainskaya as Tortula acaulon; in admixture with Tortula arenaria), LE; ibidem, Sary-Kum dune, from the dune, 17 V 2009, M. Ignatov, E. Ignatova, G. Ukrainskaya (М. Игнатов, Е. Игнатова, Г. Украинская) (det. by M. Ignatov and E. Ignatova as Tortula acaulon; with admixture of mosses), MHA 9110440; Armenia, Yerevan, semi desert with Artemisia, III 1936, A. Takhtajan (А. Тахтаджян) (det. by A. and I. Abram-ovy as Phascum cuspidatum), LE; Germany, Carlsruh (in XXIV # 5 as Phascum piliferum), LE; Romania, Transylvania (Siebenburgen), 27 II 1897, M. Peterfi (Musci Frondosi Transsilvaniae #47 as Phascum piliferum), LE; Tajikistan, Hissar Range, gorge of Varzob River, 30 km north from Dushanbe (as Stalinabad), on stony slope, 26 III 1926, P. Polyakov (П. Поляков) (det. by A. L. Abra-mova and I. I. Abramovas Phascum piliferum; in admixture with Tortula inermis), LE; ibidem, Varzob River Basin, Kondara Gorge, in thicket of almond, 120 m a. s. l., on soil, 28 II 1966, U. Mamatkulov (У. Маматкулов) (as Phascum cuspidatum), LE.
Plate IX. Tortula acaulon morphotype with mostly simple papillae ("simple papillae morphotype"). 1-3, 23 — capsules; 4, 7, 9, 10, 20, 22, 24 — stem leaves; 5 — marginal papillae; 6, 12, 18, 19, 25-27 — leaf cells; 8 — leaf margin; 11, 21 — spores; 13, 14 — leaf cross sections; 15 — perigonial leaf with antheridia; 16 — perigonial leaf; 17 —tip of leaf. Scale bars: 1-4, 7, 9, 10, 15, 16, 20, 22, 24 — 1 mm; 5, 6, 12, 18, 19, 25-27 — 20 |m; 8, 11, 21 — 40 |im; 13, 14 — 80 |m; 17 — 300 |m. 1-12 — from 7 V 2009, G. Ukrainskaya, LE; 13, 14, 18, 19 — from 4 IV 1966, K. Karczmarz, LE; 15-17, 20-27 — from 15 VIII 1988,
T. V. Sidorova, LE.
Plate X. Tortula acaulon morphotype with mostly branched papillae ("branched papillae morphotype")
1, 2 — capsules; 3 — calyptra; 4-9, 15 — stem leaves; 10, 11 — leaf cross sections; 12 — spora; 13, 14 — tip of leaves; 16-18, 22, 24 — leaf lamina cells with papillae: branched papillae in upper part of leaf (16, 22, 25), simple and 2-divided in the middle (17), simple papillae in the lower part (18); 19, 20 — costa cells with papillae: branched papillae in the middle (19), simple and 2-divided in the lower part of costa (20); 21, 23, 25 — branched papillae: C-shaped with auricles (21, 23), divided into
2-4(5) lobes (25). Scale bars: 1-9, 15 — 1 mm; 10, 11 — 80 |m; 12 — 40 |m; 13, 14 — 300 |m; 16-20, 22 — 25 |m; 21, 23 — 8 |m; 24, 25 — 20 |m. 1-11, 13-21, 24, 25 — from 27 II 1897, M. Petefi, LE; 12, 22, 23 — from 17 V 2009, G. Ukrainskaya, LE 14100.
Besides differentiation of T. irinae from T. acaulon it is necessary to discuss its distinctions from some cleistocarpous species of the family Pottiaceae Hampe with ventral outgrowths. Tortula irinae is similar to cleistocarpous species of the genus Ptery-goneurum in ventral outgrowths of costa, but differs in outgrowths presented as a pad of large, elongated parenchymatous cells (vs. as low lamellae of several rows of small cells in height in P. kozlovii and high lamellae in P. sibiricum). By ventral outgrowths of costa presented as elongated parenchymatous cells it is also similar to some species of the genus Microbryum, from which it differs in leaf lamina and costa cells with several branched papillae [vs. long-mammillose-papillose leaf lamina and costa cells with one branched papilla in M. vlassovii and long-mammillose leaf lamina and costa cells with one (below (1)2-4) mammilla in M. lydiae].
Acknowledgments. I thank bryologists of the Komarov Botanical Institute, St. Petersburg for their friendly and warm attitude to my work with herbarium material in the laboratory of Lichenology and Bryology. I am grateful to the curator of LE bryophyte herbarium Dr. Olga M. Afonina and Dr. Irina V. Czernyadjeva as well as the curator of MHA bryophyte herbarium Dr. Michael S. Ignatov for the opportunity to work freely with the herbarium material and for providing the necessary bryological literature. I am very grateful to Dr. Alexey D. Potemkin for organizing the workplace and necessary equipment, valuable advice, critical comments and his friendly help during my work on the manuscript. I thank the Director of the Komarov Botanical Institute, St. Petersburg, Dr. Dmitry V. Geltman for permission to work in the herbarium the necessary time to complete this manuscript. I thank the reviewer for the positive feedback of the manuscript and for valuable comments on research methodology.
References
Ignatov M. S., Ignatova E. A. 2003. Flora mkhov sredney chasti Rossii. T. 1. Sphagnaceae — Hedwigia-ceae [Moss flora of the Middle European Russia.Vol. 1. Sphagnaceae — Hedwigiaceae]. Moscow: 1-608. (In Russ.).
Otnyukova T. N. 2020. New cleistocarpous species of the genus Pterygoneurum (Pottiaceae, Bryo-phyta) from the steppe slopes of Siberia (Russia). Novosti sistematiki nizshikh rastenii 54(1): 251-260. https://doi.org/10.31111/nsnr/2020.54.1.251 Savicz-Ljubiskaja L. I., Smirnova Z. N. 1970. Opredelitel' listostebelnykh mkhov SSSR. Verkhoplodnye
mkhi [Handbook of mosses of the USSR. Acrocarpous mosses]. Leningrad: 826 p. (In Russ.). Zander R. H.1989. Seven new genera in Pottiaceae (Musci) and a lectotype for Syntrichia. Phytologia 65: 424-436.
Zander R. H. 1993. Genera of the Pottiaceae: mosses of harsh environments. Bulletin of the Buffalo Society of Natural Sciences 32: 1-378.
Zander R. H. M., Eckel P. M. 2007. 26. Tortula Hedwig. Flora of North America. Vol. 27. Bryophytes: Mosses, part 1. New York; Oxford: 586-603.
Литература
[Ignatov, Ignatova] Игнатов M. С., Игнатова Е. А. 2003. Флора мхов средней части России. T. 1. Sphagnaceae — Hedwigiaceae. M.: 1-608.
Otnyukova T. N. 2020. New cleistocarpous species of the genus Pterygoneurum (Pottiaceae, Bryo-phyta) from the steppe slopes of Siberia (Russia). Novosti sistematiki nizshikh rastenii 54(1): 251-260. https://doi.org/10.31111/nsnr/2020.54.1.251
[Savicz-Ljubitzkaja, Smirnova] Савич-Любицкая Л. И., Смирнова З. H. 1970. Определитель листостебельных мхов СССР. Верхоплодные мхи. Л.: 826 c.
Zander R. H.1989. Seven new genera in Pottiaceae (Musci) and a lectotype for Syntrichia. Phytologia 65: 424-436.
Zander R. H. 1993. Genera of the Pottiaceae: mosses of harsh environments. Bulletin of the Buffalo Society of Natural Sciences 32: 1-378.
Zander R. H. M., Eckel P. M. 2007. 26. Tortula Hedwig. Flora of North America. Vol. 27. Bryophytes: Mosses, part 1. New York; Oxford: 586-603.
