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Phytotaxa 299 (2): 234–235
http://www.mapress.com/j/pt/
Copyright © 2017 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
234 Accepted by Matt von Konrat: 18 Jan. 2017; published: 21 Mar. 2017
https://doi.org/10.11646/phytotaxa.299.2.7
Schistidium relictum (Grimmiaceae, Bryophyta), a new moss species from Northwest
North America and Siberia
TERRY T. MCINTOSH1, HANS H. BLOM2, OXANA I. KUZNETSOVA3 & ELENA A. IGNATOVA4
1Beaty Biodiversity Museum, University of British Columbia, 3529-6270 University Boulevard, Vancouver, British Columbia, CANADA
V6T 1Z4
2Norwegian Institute of Bioeconomy Research, Fanaflaten 4, N-5244, NORWAY
3Tsytsin Main Moscow Botanical Garden of Academy of Sciences, 127276 Moscow Botanical St., Moscow, RUSSIA
4Moscow State University, Faculty of Biology, Vorob’ovy gory 1-12, Moscow 119991, RUSSIA
Abstract
Schistidium relictum is described as a new northwest North American and Siberian species of moss. Important distinguishing
characters include dull, nearly black plants, with stems densely and evenly foliated, weakly spreading leaves that usually
lack awns, and the mostly 1-stratose distal leaf laminae with 2(–3) -stratose margins. The species has a remarkable disjunct
distribution pattern with most of the sites where it has been found having been unglaciated during the Pleistocene glaciations.
It is restricted to areas with occurrence of calcareous bedrock, especially limestones. It appears to be rather isolated geneti-
cally based on molecular studies of total ITS. It is sister to the large clade, ‘Apocarpum’, which consists of species which
probably embody its closest known extant relatives.
Key words: biogeography, internal transcribed spacer (ITS), mosses, phylogeny, Pleistocene glaciations, Schistidium
Introduction
Historically, Schistidium Bruch & Schimper (1845: 93) has been a difficult genus to resolve taxonomically (Blom
1996, McIntosh 2007, McIntosh et al. 2015). Past floristic treatments have varied from region to region, and names
that have been applied to specimens at both the species and varietal levels have been as inconsistent as the characters
used to differentiate the taxa (McIntosh 2007). However, initiated by the landmark treatment of the S. apocarpum
(Hedwig) Bruch & Schimper (1845: 99) complex in Norway and Sweden (Blom 1996), numerous Schistidium species
have been recently recognized (e.g., Blom 1996, 1998, Blom and Darigo 2009, Blom et al. 2016, McIntosh et al.
2015). The majority of these would have once been considered within the broad and uncomfortable species concept of
S. apocarpum, although a few were lodged within other broadly defined species, such as S. rivulare (Bridel) Podpêra
(1911, 207) and S. confertum (Funck) Bruch & Schimper (1845: 99). Discussions in other works have helped clarify
confusion regarding key characters useful in species recognition and include Deguchi (1979), Blom (1998), and
McIntosh (2007). A great amount of genetic research has also contributed to the understanding of species concepts
as well as species relationships within Schistidium (e.g., Goryunov et al. 2007, Ignatova et al. 2009, Hofbauer et al.
2016). As a result, although more species will be described in the coming years, Schistidium is currently comprised of
about 130 species.
The name Schistidium relictum has been used previously by H. H. Blom to name numerous herbarium collections
during his earlier research (Blom 1996). Also, it was published as a provisional name by Ignatova et al. (2009) in a key
to the Russian species of Schistidium. Because this species and its name have not, to date, been described, this name
has been considered a ‘manuscript name’. This article describes S. relictum, thus validating that name.
SCHISTIDIUM RELICTUM Phytotaxa 299 (2) © 2017 Magnolia Press • 235
Materials and Methods
Two of the authors, T. T. McIntosh and E. A. Ignatova, examined the Schistidium relictum specimens available in their
regional herbaria, whereas H. H. Blom, both during his earlier work (Blom 1998) and recently, examined the great
majority of the specimens listed below. In all cases, microscopic examination involved describing and measuring the
characters listed in the following description.
Voucher specimens of Schistidium relictum are deposited in many herbaria, i.e., University of Alaska (ALA),
University of Alberta (ALTA), National Herbarium of Canada (CANM), University of Michigan (MICH), Moscow
University (MW), Tsitsin’ Main Botanical Garden (MHA), University of Newfoundland (NFLD), New York Botanical
Garden (NY), Norwegian University of Science and Technology (TRH), University of British Columbia (UBC), and
University of California (UC).
The distribution map (Fig. 2) was made in ArcMap using a Fuller projection.
For the phylogenetic research, ITS1-5.8SrRNA-ITS2 sequences were obtained from four specimens collected
recently in northern Siberia. Specimen data and Genbank accession numbers are listed in Appendix 1 (results shown
in Fig. 3). Extractions of DNA and laboratory protocols were essentially the same as in Gardiner et al. (2005), using
White et al. (1990) primers L and B. Amplified DNA fragments were sequenced in the Center of Collective Use
“Genom” in Moscow. New sequences were included in the alignment used for broader analyses (Goryunov et al.
2007; Ignatova et al. 2010; Milyutina et al. 2010), and aligned manually in Bioedit (Hall 1999). Trees were rooted on
Schistidium sordidum following a previous analysis, where this species appeared basally in an analysis of chloroplast
DNA (Ignatova et al. 2009). Analyses were conducted under a Bayesian Markov Chain Monte Carlo approach using
MrBayes v.3.1.2 (Huelsenbeck et al. 2001) with 5.8S analyzed with HKY+I model, and ITS1 and ITS2 with a GRT+I+G
model, the model selection by ModelgeneratorV.85 (Keane et al. 2006). Three parallel runs were completed, each with
five chains and 10,000,000 generations (25% burnin), with trees sampled every 1000 generations, a temp parameter
value of 0.015.
Taxonomic Treatment
Schistidium relictum T.T. McIntosh, H.H. Blom, and E. A. Ignatova sp. nov., FIG. 1.
Diagnosis: Schistidium relictum is a distinctive species, readily separated from other species of Schistidium by a number of gametophytic
characters including dull, nearly black plants, with stems densely and evenly foliated, weakly spreading leaves that usually lack
awns, the mostly 1-stratose distal leaf laminae with 2(–3) -stratose margins, a papillose distal abaxial surface of the costa, and a
differentiated alar group of enlarged subquadrate cells delimited from adjacent cells. Also, its strongly differentiated perichaetial
leaves are much larger than adjacent stem leaves, often hiding the sporophyte in lateral view. Also, its peristome teeth are long and
curved.
Type:—CANADA. British Columbia: Mile 445, Alaska Highway, outcrop above creek, 59°05´N 125°49´E, 886 m, 26 August 1967, F.
Boas 67071 (holotype UBC! B106018, isotype TRH!).
Plants medium-sized to large, dull, nearly black or dark-brown, new growth yellow-brown, forming compact or loose
mats, tufts, or cushions, sometimes partially buried in sand or silt. Stems (1.0–)1.5–2.5(–3.5) cm, dark, upper stems
simple or branched (branching is less common in North American plants), usually somewhat curved, occasionally
straight, arching-horizontal, ascending, or erect, densely and evenly foliate and somewhat ‘columnar-like’, with broad
central strands. Stem leaves (1.2–) 1.4–1.8 (–2.0) mm long, 0.5–0.7(–1.1) mm wide, ovate-lanceolate, occasionally
lanceolate or ovate-triangular, straight or slightly secund, sometimes in ± spiral rows, sharply keeled distally, broadly
concave proximally, most leaves darker in the distal 1/2, when dry erect-appressed and sometimes with recurved
apices, when wet patent to erecto-patent and recurved, 1-stratose, not decurrent; margins weakly recurved from base
to about 2/3 (–4/5) of the leaf, smooth or papillose(-denticulate) distally, especially evident in young leaves, 2(–3)
stratose distally in 1 or 2 rows; apices acute, rarely somewhat rounded; awns usually absent, when present short, to 0.1
mm, hyaline, terete, weakly denticulate or spinulose, usually not decurrent; costae strong, dark-coloured throughout,
percurrent, dorsal surface often papillose distally, especially in young leaves, strongly bulging on abaxial surface for
much of leaf length, in transverse section rounded near apex, broadened-elliptic, asymmetric and unevenly bulging
MCINTOSH ET AL.
236 • Phytotaxa 299 (2) © 2017 Magnolia Press
medially and proximally, 2–3 cells thick; distal laminal cells (6–)8–9 µm wide, irregular in shape, short-rectangular
or more or less isodiametric, occasionally oblate, walls moderately thick to thick, not sinuose, with small corner
thickenings, smooth or with scattered, low papillae on the dorsal surface; medial laminal cells (8–)9–13 µm wide,
short-rectangular, walls moderately thick, sinuose; basal cells 12–16 µm wide, rectangular or short-rectangular, thick-
walled, weakly sinuose; alar cells differentiated as an area of enlarged and clear cells that are more or less isodiametric,
short-rectangular, or, occasionally, rectangular especially along the margins, the alar region is 1–5 rows wide and
3–6 cells high (but smaller and less defined in North American material). Perichaetial leaves paler than stem leaves,
(2.1–)2.3–3.2 mm, broadly ovate or oblong with a triangular distal portion, erect, broadly acute, often with a short
awn. Sexual condition gonioautoicous. Sporophytes common, immersed in perichaetial leaves or about 1/8 to 1/3
exposed. Seta (0.15–)0.3(–0.4) mm. Capsule urn reddish-brown to dark brown, short-cylindrical, (0.7–)0.8–1.0(–1.2)
mm, sometimes slightly narrowed towards the mouth, shiny, occasionally striolate when dry; exothecial cells short-
elongate or ± isodiametric, usually irregular in shape, sometimes 5 sided, sometimes oblate, rarely in lines, thin-walled
or slightly and unevenly thickened, walls often curved; rim the same colour as capsule wall or darker and redder.
Stomata (0–)2–6 at base of capsule. Peristome erect or patent, reflexed when dry, curved, not twisted along axis, 250–
450 µm, reddish-brown, usually densely papillose with very short papillae, usually perforated with slits, sometimes
with small terminal prongs. Operculum usually short-conic, rostrate, rostrum straight or oblique. Spores yellowish,
9–12 (–14) µm, finely granulose.
Additional specimens examined:—CANADA. British Columbia: Summit Lake, 3 mi beyond N end, 58°35´N
124°40 E, Schofield 66179 (UBC); Mt. St. George, 5300–5900 ft., 58°37´N 124°43´E, Vitt 11240 (ALTA); Wokkpash Lake,
SW shore, Mt. Roosevelt-Churchill Peak area, 58°45´N 124°85´E, Schofield, Vitt & Horton 66451 (ALTA), 66619 (ALTA,
UBC); Muncho Lake area, Alaska Highway post 444.9 at Peterson Creek Bridge No 4, 58°52´N 125°49´E, Vitt & Andrus
3023 (ALTA); Rocky Mts., Haworth Lake, 3800 ft., 58°47´N 125°07´E, Vitt 20096 (ALTA, with Schistidium trichodon);
Yukon: Northern Ogilvie Mts., Nahoni Range, 65°37´N 139°06´E, Vitt 12963 (ALTA, with Schistidium boreale); Northern
Ogilvie Mts., Nahoni Range, 4600–4700 ft., 65°37´N 139°03´E, Vitt 13073 (ALTA); Northern Ogilvie Mts., Nahoni
Range, 65°38´N 139°03´E, Vitt 13378 (ALTA, NFLD); Northern Ogilvie Mts., Nahoni Range, 3900–4100 ft.,65°37´N
139°02´E, , Vitt 13130, 13156 (ALTA); mountain S of Mt. Klotz, at headwaters of Ogilvie River, 65°20´N 140°06´E,
Vitt 7482 (ALTA); Mackenzie Mts., Bonnet Plume Range, Wrightii Lake, ca 7.5. mi SE of Fairchild, 3300–4400 ft.,
65°54´N 133°33´E, Horton 6701 (ALTA), Vitt 16977 (ALTA); Bonnet Plume Range, Riparium Lake, 4800–5200 ft.,
64°46´N 133°23´E, Horton 6559 (ALTA); South Richardson Mts., Doll Creek area, 66°05´N 135°48´E, Ritchie 7045
(CANM); Peel River Watershed, Mt. Reception, site 36, 65°37´N 135°30´E, Bird & Benson 30496 (CANM); N.W.T.:
Inuvik, SE of Airport Lake along stream, Packer & Lemay 205 (ALTA, with Grimmia teretinervis); Mackenzie distr.,
Nahanni Range, just N of peak at 3598 ft., 61°43´N 123°20´E, Vitt 20290 (ALTA, NY); Nahanni Range, 62°13´N
123°22´E, Horton 10403 (ALTA); Mackenzie distr., Ram Range, 61°43´N 123°53´E, Vitt 20390 (ALTA); Campbell
Hills, 68°14´N 133°15´E, Scotter 28358 (ALA, + Schistidium submuticum subsp. arcticum); Gibson Ridge, Chick
Lake, 1300 ft., 65°49´N 128°15´E, Gubbe & Burr 616A (ALTA); South Nahanni River Natl. Park, Deadman Valley,
61°19´N 124°35´E, Steere 76-325 (NY); Kraus Hot Springs, 61°15´N 124°03´E, Steere 76-291 (NY), Scotter 22530
(NY); Virginia Falls, 61°38´N 125°42´E, Scotter 22349, 22259, 22260 (NY); Nunavit: Devon Island, 75°40´N 84°40´E,
Vitt 5354 (ALTA); U.S.A. Alaska. Central Yukon River district, White Mts., limestone ridge, Gjærevoll 18.VII.1953 (TRH),
Gjærevoll 28.VII.1953 (TRH); Chandalar Quad., Brooks Range, Sukapak Mt., Spatt 631 (ALTA); base of Sukapak Mt.,
610 m, 67°36´N 149°45´E Murray 77-121 (ALA); Mountains NW of Walker Line, upper Kopuk River, above W end of
Walker Lake, 67°10´N 154°30´E, Jordal 4041 (MICH); Survey Pass Quad., vicinity of confluence of Alatna and Nahtuk
rivers, 1200 ft., 67°25´N 153°43´E Murray 73-42 (ALA); vicinity of Ambresvajun Lake (= Last Lake), 1159 m, 68°38´N
143°41´E, Batten 75-43c (ALA); Endicott Mts., Arrigetch Creek valley, 3400 ft., 67°26´N 154°05´E, Cooper CB-161,
4500 ft., 67°26´N 154°00´E, Cooper CB-165 (ALA); Okpilak Lake vicinity, 69°25´N 144°03´E, Murray 8625 (ALA);
Toolik Lake, Wallace 18.VII.1976 (ALA); Arctic Village, White mountain W of Chandalar River, 68°08´N 145°32´E,
Steere & Iwatsuki 74-293 (NY); RUSSIA (ASIA): West Siberian Arctic, Yamalo-Nenetzky Autonomous District, Yunto
Lake, 67°40´N, 68°00´E, 10.VIII.1993, Czernyadjeva 58 (LE, MW); Krasnoyarsk Territory: Taimyrsky Autonomous
District, State Biosphere Natural Reserve Taimyrsky, Medvezh’ya (Bear’s) River, slope of a limestone ridge along the
right bank of Kotuy River, 70°58´N, 102°49´E, Fedosov 05-672 & 05-718 (MW); Taimyrsky Autonomous District,
right bank of Maimecha River 14 km downstream of Chopko Creek mouth, 70°50´N, 100°57´E, 30.VI.2009, Fedosov
09-46 (MW); Republic Sakha/Yakutia: Bulun District, lower course of Lena River, Kharaulakh Mt. Ridge, Ogonnior-
Yurege Creek, 170 m, 71°25´N, 127°25´E, 09.VII.2006, Pisarenko op04672 (MW); Tompo District, Sette Daban
Mt. Ridge, right bank of Segenyakh (Rosomakha, or Wolverine) Creek upstream Magadan Hwy, 470 m. 63°02´N,
137°57´E, 17.VII.2015, Ignatov & Ignatova 15-538 (MHA, MW).
SCHISTIDIUM RELICTUM Phytotaxa 299 (2) © 2017 Magnolia Press • 237
FIGURE 1. Schistidium relictum. A, F. Habit. B–C. Sporophytes, with and without operculum. D. Stem section. E. Cells of stem leaf
apical portion with hyaline tip. G. Peristome tooth. H. Upper laminal cells. I. Median laminal cells. J. Basal laminal cells. K. Perichaetial
leaf. L–N. Cauline leaves. O–R. Leaf transverse sections. S. Exothecial cells. Scale bars: 5 mm for F; 2 mm for A–C; 1 mm for K–N; 100
µm for D, G, O–R; 50 µm for E, H, I–J, S.
MCINTOSH ET AL.
238 • Phytotaxa 299 (2) © 2017 Magnolia Press
FIGURE 2. Global distribution of Schistidium relictum.
Etymology:—The epithet relictum refers to a distribution pattern that suggests that S. relictum survived large
parts of the Pleistocene glaciations in situ as a relict.
Distribution and Habitat:—Schistidium relictum is known from northwest North America and northern parts
of Siberia in Asian Russia (Fig. 2). Its elevation range is from ca 100–470 m in Siberia and ca 50–1700 m in North
America. The species has a remarkable disjunct distribution pattern with most of the sites where it has been found
having been unglaciated during long periods of the Pleistocene glaciations (e.g., Svendsen et al. 2004). We hypothesise
that the species persisted in ice-free refugia north of the continental ice sheets during the Pleistocene, the huge Beringian
refugium being the most important one (e.g., Hultén 1937, Schuster 1983). Its present distribution pattern probably
indicates a low degree of migration in the Holocene following the retreat of the inland ice sheets.
Schistidium relictum is restricted to areas with occurrence of calcareous bedrock, especially limestones. It has been
collected on dry ledges, walls, and boulders but also on periodically irrigated rocks on river banks, and occasionally on
mineral soil. According to specimen labels and field data, there is a rather large variation in habitats from windswept
ridges with little snow cover to adjacent to snow beds to moist sheltered sites in creeks and along rivers. Based on species
admixtures in herbarium specimens, it grows with other Grimmiaceous calciphytes including Schistidium submuticum
subsp. arcticum H.H.Blom, S. frisvollianum H.H.Blom, S. boreale Poelt, and Grimmia teretinervis Limpr.
The distribution of Schistidium relictum in North American resembles that of Andreaeobryum macrosporum
Steere & B.M. Murray, which also occurs along the ice free corridor between the Laurentide and Cordilleran ice sheets
in the Yukon-Northwest Territoiry-northern British Columbia area (Pedersen et al. 2016). Further, Schistidium relictum
was recently discovered in Yakutia, east Russia (Ivanova et al. 2016), and, interestingly, was collected at this site along
the same creek as Andreaeobryum macrosporum from 500 to 1100 m elevation. Both species are likely not rare in the
mountains of this area.
Differentiation and relationships:—Schistidium relictum is a distinctive species only likely to be confused with
other blackish Schistidium species, in particular S. andreaeopsis and S. boreale. Schistidium andreaeopsis is a much
larger plant with longer leaves (2.0–2.7 mm vs., on average, 1.4–1.8 mm) and wider distal leaf cells (10–12 µm vs.
8–9 µm). Also, its leaf cells are strongly sinuose to nodulose almost throughout and possess characteristic reddish
walls, whereas they are esinuose or slightly sinuose in upper part of leaf and lack the reddish wall coloration in S.
relictum. Sporophytes are unknown in S. andreaeopsis. The gametophyte of S. boreale is similar to S. relictum, but its
leaf laminal cells are distinctly more densely and highly papillose and cells walls often reddish or orange. Also, the
peristome teeth of S. boreale are shorter than those of S. relictum (220–330 µm vs. 250–450 µm).
SCHISTIDIUM RELICTUM Phytotaxa 299 (2) © 2017 Magnolia Press • 239
FIGURE 3. Illustration of genetic relationships of Schistidium relictum with selected species and species groups of Schistidium based on
molecular studies of total ITS.
MCINTOSH ET AL.
240 • Phytotaxa 299 (2) © 2017 Magnolia Press
Schistidium relictum appears to be rather isolated genetically based on molecular studies of total ITS (Fig. 3). It
is sister to the large clade, ‘Apocarpum’, which consists of species which probably embody its closest known extant
relatives. Schistidium relictum shares the character states of long peristome teeth, papillose leaf costae, and denticulate
upper leaf margins with several of these species. The tree in Figure 3 includes a basal paraphyletic clade formed by S.
pulchrum, S. grandirete, S. sibiricum, and S. frisvollianum, and the main clade (PP=0.99) that includes all remaining
species. The latter is subdivided into two clades, which, however, have low support. In contrast, high support is present
for clades that correspond to the various sections, established by Blom (1996) based on morphological study, and
clades from the previous broader analyses: Atrofuscum-clade (PP=1.00), Frigidum-clade (PP=0.96), Confertum-clade
(PP=0.79, but P=0.99 for the main volume of this clade excluding S. tenerum), Apocarpum-clade (PP=0.99), and, in
addition, the clade formed by four specimens of S. relictum. The latter retains the sister position to the Apocarpum-
clade, and, even though with only moderate support (PP=0.75), indicates the isolated position of S. relictum.
Acknowledgments
Sincere thanks to the curators of the North American herbaria listed above for arranging loans to H.H. Blom during
his previous studies of specimens of Schistidium, and to Line Walberg Larsen, Ås, for preparing the distribution
map. Thanks also to Catherine La Farge for assisting in gathering herbarium data at the University of Alberta. The
phylogenetic work was supported by Project 14-50-00029 (Scientific basis of the national biobank—depository of the
living systems) funded by a grant from the Russian Science Foundation (RSF).
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242 • Phytotaxa 299 (2) © 2017 Magnolia Press
APPENDIX 1
Schistidium abrupticostatum_Taimyr_188 HM053930; S. abrupticostatum_Severnaya_Zemlya_194 HM053928; S.
agassizii_Murmansk_158 HM053878; S. agassizii_Taimyr_160 HM053879; S. andreaeopsis_Taimyr_85 HM053881;
S. andreaeopsis_Canada_138 HM053882; S. apocarpum_Caucasus_129 DQ822033, HM031074; S. apocarpum_
United_Kingdom_126 HM031076; S. boreale_Urals_127 DQ822024, HM031069; S. boreale_Sweden_80 HM053890;
S. canadense_Karelia_130 DQ822016, HM031067; S. canadense_USA_Maine_103 HM053917; S. confertum_
Austria_169 HM053891; S. confertum_Austria_33 JF262179; S. crassipilum_Poland_21 DQ822020, HM031073; S.
dupretii_Austria_167 HM053894; S. echinatum_Austria_31 HQ890509; S. echinatum_Caucasus_304 HQ890508; S.
elegantulum_Caucasus_32 DQ822022, HM031071; S. flaccidum_Caucasus_177 HM053896; S. flaccidum_Caucasus_
171 HM053899; S. frigidum_Taimyr_109 HM053905; S. frigidum_Taimyr_146 HM053907; S. frisvollianum_Taimyr_
150 HM053908; S. frisvollianum_Taimyr_172 HM053909; S. grandirete_Taimyr_87 HM053910; S. grandirete_
Severnaya_Zemlya_136 HM053911; S. lancifolium_Sakhalin_206 HM053912; S. lancifolium_USA_Maine_193
HM053917; S. marginale_Austria_86 HM053919; S. marginale_Austria_84 HM053920; S. maritimum_Murmansk_
165 HM053922; S. maritimum_subsp_piliferum_Norway_170 HM053923; S. obscurum_Taimyr_107 HM053900;
S. obscurum_Austria_236 HQ890519; S. obscurum_Spitsbergen_140 HM053903; S. papillosum_Caucasus_13_124
DQ822012, HM031061; S. pulchrum_Urals_34_180 DQ822030, HM031053; S. pulchrum_Taimyr_215 DQ822031,
HM031050; S. relictum_Taimyr_t82 KX522944; S. relictum_Yakutia_1154 KX522946; S. relictum_Yakutia_1125
KX522947; S. relictum_Yakutia_1153 KX522945; S. rivulare_Altai_198 HM053934; S. rivulare_Kuril_Islands_197
HM053936; S. robustum_Sweden_148 HM053938; S. scandicum_Urals_133 DQ822026, HM031054; S. scandicum_
Sweden_179 DQ822027, HM031059; S. sibiricum_Buryatia_79 HM053883; S. sibiricum_Chita_69 HM053884; S.
sordidum_Taimyr_182 HM053942; S. sordidum_Yakutia_183 HM053943; S. strictum_Norway_108 HM053944;
S. subflaccidum_Austria_89 HM053945; S. subflaccidum_Caucasus_74 HM053946; S. subjulaceum_Altai_216_
HQ890522; S. subjulaceum_Buryatia_118 HM053947; S. submuticum_Urals_128 DQ822011, HM031055; S.
submuticum_Urals_123 DQ822010, HM031056; S. succulentum_Taimyr_106 HM053875; S. succulentum_Caucasus_
305 JF280964; S. tenerum_Canada_142 HM053951; S. tenerum_Chukotka_99 HM053952; S. tenuinerve_Altai_306
HQ890525; S. tenuinerve_Sakhalin_233 HQ890524; S. trichodon_var_nutans_Austria_71 HM053953; S. trichodon_
var_nutans_Caucasus_114 HM053954; S. viride_USA_Maryland_102 HM053958; S. viride_USA_Missouri_104
HM053957.
