ON SCIURO-HYPNUM OEDIPODIUM AND S. CURTUM - Arctoa

Arctoa (2007) 16: 47-61 

ON SCIURO-HYPNUM OEDIPODIUM AND S. CURTUM 

(BRACHYTHECIACEAE, BRYOPHYTA) 

О SCIURO-HYPNUM OEDIPODIUM И S. CURTUM 

(BRACHYTHECIACEAE, BRYOPHYTA) 

MICHAEL S. IGNATOV 1 & IRINA A. MILYUTINA 2 

МИХАИЛ С. ИГНАТОВ 1 & ИРИНА А. МИЛЮТИНА 2 

INTRODUCTION 

The complex of species around Scuiro-hypnum 

(formerly Brachythecium) starkei is known 

to be difficult, causing considerable incongruence 

of species concepts of different authors during the 

XIX and most of the XX century. 

A detailed overview of this history was presented 

by Piippo (1983), who also selected the neotype 

of Hypnum starkei Brid. and lectotypes of H. 

oedipodium Mitt. and H. curtum Lindb., i.e. the 

taxa in which the indefinite concepts were the main 

source of taxonomic and nomenclatural confusions. 

According to Piippo (1983), this complex includes 

two species in the Northern Hemisphere, 

Abstract 

Sciuro-hypnum curtum has been synonymized with S. oedipodium, but the present revision 

reveals that they are two distinct species although morphological variation is very broad in both of 

them. However, molecular delimitation of these two species (nrITS) is much more sharp. Sciurohypnum 

curtum is a widespread species in boreal forests of Eurasia, and occurs also in North 

America mainly in its eastern part, with solitary localities in the West. Sciuro-hypnum oedipodium 

is widespread in western North America, totally absent in the eastern part of this continent, and is 

known in Russia in Caucasus and in Chukotka. Brachythecium holzingeri is synonymized with S. 

oedipodium. 

Резюме 

Sciuro-hypnum curtum был отнесен с синонимы к S. oedipodium и последние 25 лет не 

считался самостоятельным видом, но настоящее исследование установило, что эти два 

вида довольно далеки друг от друга, несмотря на то, что из-за сильной вариабельности 

некоторые их морфотипы внешне очень сходны. Молекулярные данные по последовательностям 

ITS подтверждают их весьма значительные отличия, при том что каждый из них 

константен на протяжении всего ареала. Sciuro-hypnum curtum широко распространен в 

таежных лесах Евразии, встречается на востоке Северной Америки и в единичных точках 

на ее западе. Sciuro-hypnum oedipodium широко распространен на западе Северной 

Америки, отсутствует на востоке этого континента, а также выявлен в России на Кавказе 

и на Чукотке. Brachythecium holzingeri отнесен в синонимы к S. oedipodium. 

Brachythecium oedipodium and B. starkei (this 

group of Brachythecium was later transferred to 

Sciuro-hypnum by Ignatov & Huttunen (2002)). 

Both these species are widespread in northern 

Europe occurring also in Asia and North America. 

The concept of Piippo received wide acceptance 

and was followed in checklists of North 

America (Anderson et al., 1990), China (Redfearn 

et al., 1996), Europe (Hill et al., 2006), East Europe 

and North Asia (Ignatov,Afonina, Ignatova 

et al., 2006). 

The work on the treatment of Brachytheciaceae 

for the Flora of North America, however, 

revealed certain difficulties in the application of 

1 — Main Botanical Garden of Russian Academy of Sciences, Botanicheskaya, 4, Moscow 127276 Russia – 

Россия 27276 Москва, Ботаническая, 4, Главный ботанический сад РАН 

2 — A.N.Belozersky' Research Institute of Physico-Chemical Biology, Moscow State University, Moscow 119991 

Russia – НИИ Физико-химической биологии им. А.Н. Белозерского, МГУ, Москва 119991 [fax: 7-(095)- 

939-31-81; e-mail:tav@genebee.msu.ru]

48 MICHAEL S. IGNATOV & IRINA A. MILYUTINA 

Piippo’s (1983) concept. The species identical to 

widespread European ‘oedipodium’ sensu Piippo 

was found to be more or less widespread in New 

England and neighboring areas in eastern North 

America (but with two localities in the Rocky 

Mountains), while the type of Hypnum oedipodium 

was from the mountains of the West. The type 

itself comprises poorly developed plants, however 

very numerous collections from the western 

states, usually kept in North American herbaria 

(e.g. in DUKE, MO, NY, US) under ‘Brachythecium 

starkei’, represent a species quite different 

from ‘oedipodium’ sensu Piippo. The type of 

Hypnum oedipodium, as it will be argued below, 

belongs to this widespread western species, and 

therefore cannot be applied to the widespread 

Eurasian and eastern North American species. The 

earliest species name for the latter species and 

also the name used for a long time in Europe is 

‘curtum’, thus this taxon has to be called: 

Sciuro-hypnum curtum (Lindb.) Ignatov, 

comb. nov. – Hypnum curtum Lindb., Musci 

Scandinavici 35. 1879 [the validation in Helsingfors 

Dagblad 1878 (342): 2. 1878 is earlier, but 

probably has to be ignored as it appeared in a 

public newspaper, not in scientific literature] – 

Brachythecium curtum (Lindb.) J. Lange & 

C.O.E.Jensen, Medd. Groenland. 3: 337. 1887. 

At the same time, Sciuro-hypnum starkei s. 

str., identical with European and Asian, occurs in 

North America, although limited in its distribution 

to the eastern provinces of Canada, occurring 

in U.S.A. only in Maine, Vermont and Wisconsin. 

MORPHOLOGICAL DIFFERENCES BETWEEN 

S. CURTUM AND S. OEDIPODIUM 

Both species are very variable, so the marginal 

phenotypes and poorly developed plants of 

these species (as well as many other Brachytheciaceae) 

are difficult or even impossible to interpret. 

The optimally developed, S. curtum (cf. Fig. 

1) is a relatively large plant with rather remotely 

arranged leaves, that are patent and usually 

somewhat twisted along their length when dry, 

resulting in a peculiar appearance that has an 

aspect of terrestrial phenotypes of Plagiothecium 

denticulatum. Branches have an apparent tendency 

to complanate foliage. Stem leaves are 

broadly ovate below and are broadest at 1/7–1/10 

of leaf length. Decurrencies are long but rather 

narrow. Mid leaf cells are linear, 60-80 μm long, 

being 7-12:1. Cells in leaf corners are enlarged, 

thin-walled and thus pellucid, gradually or rather 

abruptly delimited from laminal and basal juxtacostal 

cells. Branch leaves are usually strongly 

serrate. Seta is distinctly and invariably rough. 

Capsule is relatively long, and when young is 

strongly curved with operculum pendent to faced 

back towards the seta, or ultimately even faced 

upward, the capsule having made a curve of >270° 

[there are however exceptions when capsule is 

relatively short]. The under-developed plants have 

about the same size of stem and branches, but 

smaller and more remote leaves. 

The optimally developed plants of S. oedipodium 

(e.g. Fig. 2) are rather densely and terete 

foliate, so the plants have the appearance of 

Brachythecium or of a very large Sciuro-hypnum 

glaciale. Leaves are appressed at their bases, and 

above are either erect, or gradually reflexed. 

Leaves are ovate, but compared with S. curtum 

are not that broad and usually have broadest point 

at 1/6–1/7 of leaf length (not at 1/7–1/10 or occasionally 

even below the point of insertion in 

S. curtum). Decurrencies are long and conspicuously 

broad (this is a useful character for identification 

of smaller phenotypes). Mid-leaf cells 

are very variable even within one shoot, ranging 

from 3-4:1 to 7-10:1; an example is shown in 

Fig. 5. Cells in the leaf corner are subquadrate 

to shortly rectangular, slightly enlarged, rather 

numerous along the margin to the broadest point 

of leaf, with ‘normally’ thickened cell walls, thus 

the alar group is conspicuous, but ‘not especially 

pellucid, not especially opaque’, usually very 

gradually transforming into laminal and basal 

juxtacostal cells. Branch leaves usually serrulate, 

but variable in this character from subentire 

to quite serrate. Seta is distinctly rough, although 

in a few collections only slightly so. Capsule 

is short, inclined to horizontal, and in many 

cases looks ‘too abruptly’ hozirontal for that 

short shape; more rarely the capsule is somewhat 

elongate, but at the immature state is never 

strongly curved. The under-developed plants are 

smaller and with smaller leaves and the variation 

from large to small shoots is observed in 

many collections.

2 mm 

5 

12 

6 

Sciuro-hypnum oedipodium and S. curtum 

2 

7 

200 μm 

8 

1 

1 mm 

Fig. 1. Sciuro-hypnum curtum (Lindb.) Ignatov (from 

U.S.A., NY, Allen #27150 (MO)): 1 – seta; 2 – habit; 3, 11 

– mid-leaf cells; 4 – upper leaf cells; 5-7 – branch leaves; 8- 

10 – stem leaves; 12-13 – basal leaf cells. Scale bars: 2 mm 

for 2; 1 mm for 5-10; 200 μm for 1; 100 μm for 3-4, 11-13. 

9 

3 

13 

4 

10 

11 

100 μm 

49


2 mm 

9 

3 

2 

1 

100 μm 

12 

10 11 13 

Fig. 2. Sciuro-hypnum oedipodium (Mitt.) Ignatov & Huttunen (from U.S.A., Wyoming, 

Eckel 94081601, MO): 1 – capsule; 2-3 – habit; 4-7 – stem leaves; 8 – upper leaf cells; 9-10 – 

mid-leaf cells; 11-13 – branch leaves; 14 – basal leaf cells. Scale bars: 2 mm for 1-3; 1 mm for 

4-7; 11-13; 100 μm for 8-10, 14. 

4 

1 mm 

5 

6 7 

8 

14

1 

2 mm 

2 

Fig. 3. Sciuro-hypnum oedipodium (Mitt.) Ignatov 

& Huttunen (from the lectotype of Hypnum oedipodium 

Mitt., NY): 1-4 – habit; 5-7 – capsules; 8 – stem leaf; 9 

– mid-leaf cells; 10 – basal leaf cells. Scale bars: 2 mm 

for 1-7; 1 mm for 8; 100 μm for 9-10. 

TYPE OF HYPNUM OEDIPODIUM 

Piippo (1983) selected the lectotype of Hypnum 

oedipodium from several collections of Lyell 

from Pack River in the Northern Idaho. Two other 

collections from the same place were designated 

as syntypes, and are identical, except the collection 

from Michigan that belongs to S. curtum. 

The lectotype is correctly selected, described 

and illustrated by Piippo (1983). However the collection 

does not represent plants that are very well 

developed, thus providing problems with its interpretation. 

In Fig. 3, 8-10, one of the stem leaves 

from a small envelope where Piippo has put leaves 

used for her drawings is illustrated. Fig. 3, 10 

shows alar cells typical for plants widespread in 

the West, and leaf shape (Fig 3, 8), mid-leaf cells 

(Fig. 3, 9) and short capsules (Fig. 3, 5-7), as well 

as non-complanate foliage (Fig. 3, 1-4). These 

features demonstrate that the type of Hypnum 

oedipodium and the species widespread in moun- 


3 

4 

5 

6 

1 mm 

7 

8 

100 μm 

9 

10 

51 

tains of western North America belong to a single 

species and therefore the name ‘oedipodium’ 

should be used for it. 

WHAT IS BRACHYTHECIUM HOLZINGERI? 

Grout described this species first as a variety 

of Brachythecium collinum (Grout, 1901), then 

raised its status up to species level (Grout, 1922), 

then decided that it was not a member of Brachytheciaceae 

at all (Grout, 1928), and finally accepted 

it again as a good species of Brachythecium 

(Grout, 1934). This vacillation might look strange 

to one who has never worked on taxonomy of 

Brachytheciaceae, where many species exhibit 

‘intergradations’ ‘to all other species’. Note that 

Brachytheciastrum (Brachythecium) collinum is


1 

2 mm 

1 mm 

6 7 8 9 

Fig. 4. Sciuro-hypnum oedipodium (Mitt.) Ignatov & 

Huttunen (from holotype of Brachythecium collinum var. 

holzingeri Grout, NY): 1 – habit; 2-3 – mid-leaf cells; 4 

– basal leaf cells; 5 – upper leaf cells; 6-10 – leaves. Scale 

bars: 2 mm for 1; 1 mm for 6-10; 100 μm for 2-5. 

one of the smallest species of Brachythecium s. 

ampl., while here B. collinum var. holzingeri is 

synonymized with Sciuro-hypnum oedipodium, 

one of the largest species of Brachythcium s. ampl. 

This case demonstrates once more the extreme 

polymorphism of Sciuro-hypnum oedipodium. 

Sciuro-hypnum oedipodium (Mitt.) Ignatov 

& Huttunen, Arctoa 11: 270. 2002 [2003]. – 

Brachythecium collinum var. holzingeri Grout, 

Bryologist 4: 236. f. 2a. 1901, syn. nov. – 

Brachythecium holzingeri (Grout) Grout, Bryologist 

25: 14. 1922. 

The holotype of Brachythecium collinum var. 

holzingeri in DUKE has a printed label inside 

the envelope: ‘Mosses of North-Western Montana, 

from the vicinity of lake McDonald, Flathead 

county, N.-W. Montana. The lake in four 

miles north of Belton, a station 30 miles east of 

Kalispell, on the Great Northern Railway. Distributed 

by J.M. Holzinger. Collected by John 

M. Holzinger and James B. Blake, July 14-30, 

Fig. 5. Sciuro-hypnum oedipodium (Mitt.) Ignatov 

& Huttunen (from Colorado, MO): mid-leaf areolation 

from different leaves of one shoot, showing variation. 

Scale bar: 100 μm. 

2 

3 

10 

4 

1898, Altitude 3,500–7,000 feet.’ Handwritten 

on this label: ‘Base of Sperry glacier, July 25.’ 

There are three labels glued on the envelope: 

1) Rev. N.A.Brachytheciaceae, A.J.Grout, 

B. collinum var. holzingeri. Leaves larger and 

more gradually acuminate than in B. collinum. 

Less serrate leaf, cells broader, [unclear]. 

2) belong to the collinum group 

3) type. 

[inside there are also two other labels, one with 

illegible handwriting, second (printed): ‘North 

American Musci Perfecti issued by A.J.Grout, 

Ph.D.’, pencil handwriting of A.J.Grout ‘May 

1928, This seems to be Amblystegium Kochii forma’]. 

5 

100 μm 

100 μm

The collection is very poor, comprised mostly 

of Polytrichum with a few thin shoots of a Brachytheciaceae, 

which exhibit however the main diagnostic 

characters of S. oedipodium (Fig. 4), and 

therefore is synonymized with this species. 

The type of ‘holzingeri’ differs from B. collinum 

in the less strong margin serration and less 

thick-walled cells. 

THE OCCURRENCE OF S. OEDIPODIUM IN 

EURASIA 

The circumspription of S. oedipodium in North 

America has helped in the interpretation of several 

enigmatic specimens from Russia, that were 

kept in a folder ‘to be described’. 

Plants from middle and upper elevations of 

Caucasus (Fig. 6) could not be comfortably placed 

in S. curtum, because of too long leaves, opaque 

rather than pellucid alar cells and relatively short 

capsule. From ITS sequences it appears to be closest 

to S. reflexum, although strikingly different 

from the latter in the much bigger size of the plants 

(like large Brachythecium salebrosum or B. rutabulum). 

The ITS sequence differs from that of 

S. reflexum by just one substitution, but that is 

the same substitution by which S. reflexum differs 

from the American S. oedipodium (see below 

for details). 

In general, S. oedipodium seems to be rather 

rare in Caucasus, and so far we have failed to discover 

it in other Caucasian countries, although it 

likely occurs in Georgia. This rarity might also correspond 

to its rarity in Central European mountains. 

We found no specimens in MW and LE. At 

the same time, S. oedipodium is likely present there, 

judging from the overall distribution, and from the 

descriptions of Limpricht (1896). In addition to 

Brachythecium curtum, Limpricht also recognized 

Brachythecium strakei, and the description and illustrations 

of the latter species are in good agreement 

with the current concept of S. oedipodium. 

Sciuro-hypnum starkei, as it is understood now (cf. 

Piippo, 1983, and see also below) does not fit any 

of Limpricht’s descriptions, although according to 

the key to species ‘Brachythecium troemsoense 

(Kaur. & Arnell) Limpr.’ seems to be fairly close 

to S. starkei, as it has a percurrent costa (however 

the species description indicates a short costa). 

In addition, S. oedipodium was found in just 

one collection from Chukotka, where it could be 


53 

expected, judging from its distribution in the 

north-west of North America. The Gil’mimliveem 

Hot Springs, where it was collected, are not far 

from another hot springs with the only Asian locality 

of Plagiothecium undulatum, a species common 

in Western North America, also exhibiting 

disjunction to Europe (Afonina, 2004). 

THE CURRENTLY KNOWN DISTRIBUTION OF S. 

CURTUM AND S. OEDIPODIUM 

The main purpose of this paper is to stress the 

non-identity of S. curtum and S. oedipodium that 

was found during revision of North American and 

Russian material, thus only a few collections from 

Eurasia outside Russia were revised and included 

in the list below. Mostly just one specimen per 

country is cited, except Russia, U.S.A. and Canada, 

where specimens are cited one per state/province/territory/ 

or other administrative unit. 

SELECTED SPECIMENS EXAMINED: 

Sciuro-hypnum curtum 

EUROPE: RUSSIA: Arkhangelsk Prov.: Kargopol, 

Churakova #1238 (MHA); Bashkiria: Malyj Iremel 

Mt., 700 m, Ignatova #2/65 (MHA); Belgorod 

Prov.: Shebekino Distr., Titovka, 12.VII.2006 Maslova 

(BSU ex MHA); Bryansk Prov.: Zhukovka Distr.: 

between Zhukovka and Trosna, 23.VII.1974 Makarov 

(MHA); Chuvashia: Prisursky reserve, V.1999 Moshkovsky 

(MHA); Kaluga Prov.: Mechivino, 16.VIII.1911 

Nikolaevskaya (LE); Karelia: Kindo Cape, 29.VI.1994 

Spirina (MW); Komi Republic, 22 km WNW of Syktyvkar, 

27.VI.1994 Zheleznova (MHA); Kostroma 

Prov.: Sharya Distr., Pechenkino, 17.IX.1990 Ignatov 

(MHA); Kursk Prov.: Centralno-Czernozemny Reserve, 

13.VIII.1996 Ignatov (MHA); Leningrad Prov.: 

Komarovo 26.VII.2007 Kurbatova (LE); Lakhta, 

4.IX.1919 anonymous (MW); Mari-El Republic: 

Bolshaya Kokshaga Reserve, 15.X.1999 Bogdanov et 

al., (MHA); Moscow Prov.: Taldom Distr., Meldino, 

18.VII.1996 Ignatov & Ignatova (MHA); Murmansk 

Prov.: Ponoy River, Czernyadjeva 30.VII.1989 (LE); 

Nizhny Novgorod Prov.: Kerzhensky Reserve, 

15.X.1998 Popov #203 (MHA); Novgorod Prov.: 

Valdaj Distr., 1.VIII.1981 Morozova (MHA); Perm 

Prov.: Vishera Reserve, 500 m, 30.VI.1994 Bezgodov 

#290 (MW); Pskov Prov.: Mikhailovskoe, 17.VIII.2005 

Afonina (LE); Ryazan Province: Oksky Reserve, 

2.VI.1968 Bogdanova (MW); Smolensk Prov.: Petrakovo, 

16.VII.2004 Ignatov (MHA); Tatarstan: Volzhsko-Kamsky 

Reserve, 18.VIII.2003 Ignatov & Ignatova 

(MHA); Tula Prov.: Yulskie Zaseki, 27.VI.1971 

Kurnaev (LE); Udmurtia, Votkinsk Distr., Kostovaty, 

17.VII.2000 Munitzyna (MHA); Vladimir Prov.: Gor-


4 

3 

1 

6 

2 mm 

5 

1 mm 

100 μm 

7 8 9 

Fig. 6. Sciuro-hypnum oedipodium (Mitt.) Ignatov & Huttunen (from Russia, Caucasus, Teberda Reserve, 

2100 m alt., Ignatov & Ignatova 06-3038): 1 – habit; 2 – upper leaf cells; 3-4 – mid-leaf cells; 5-6 – stem leaves; 

7-8 – branch leaves; 9 – basal leaf cells. Scale bars: 2 mm for 1; 1 mm for 5-8; 100 μm for 2-4, 9. 

2

okhovets Distr.: 32 km NNE of Murom, Seregin #M- 

1565 (MHA); Vologda Prov.: Sokol Distr., Shorega 

Creek 22.IX.1990 Ignatov (MHA). GERMANY: Breutel, 

Flora Germ. Exs. #387 (LE, sub. Hypnum starkei). 

POLAND: Ochyra - Musci Pol. Exs. 878,.1182 (MHA, 

sub. Brachythecium oedipodium). SWEDEN: Stockholm, 

25.II.1995 Ignatov (MHA). BELARUS: Grodno 

Prov., Seregin et al., #M-1673 (MHA); UKRAINA: Kiev, 

19.VII.1986 Virchenko (MHA); ESTONIA: Keeri-Karijдrve 

Nature Reserve, 31.VIII.2007 Leis (MHA). 

ASIA: RUSSIA: Altai Republic: Bolshoye 

Istyube Creek, 470 m, Ignatov 18/44, 18/9 (MHA); 

Altaisky Territory: Barnaul, 21.VII.2004 Ignatov 

(MHA); Amurskaya Prov.: Zeysky Reserve, 5.VIII. 

1980 L.I. Abramova (MHA ex IRK); Buryatia: Tompuda, 

1500 m, 18.VIII.1956 Bardunov (MHA ex IRK); 

Evenkiya: Kochumdek Creek, 20.VIII.1991 Shcherbina 

(MW); Khakassia: Shirisky Distr.: Yarovoe, 

12.VIII.1961 Kamenetskaya (MHA ex IRK); Khanty-Mansijsk 

Autonomous Area: Irtysh, 120 km 

downstream from Tobolsk, 8.VIII.1973 Iljina & Kobeleva 

(MHA ex IRK); Krasnoyarsk Territory: 

Boguchansk Distr., Chuny, 4.VIII.1988 Kustova 

(MHA ex IRK); Kulumys Range, 17.VI.1968 Bardunov 

(MHA ex IRK); Tomsk Prov.: Kaftannikovo, 

11.VIII.1913 (LE); Primorsky Territory: Shkotovo 

Distr. 15.IX.947 Rosenberg (LE); Tyumen Prov.: 

Berezovsky Distr. 4.X.1950 (LE); Yakutia: Lensky 

Distr., Pilka River 25.VII.1999 Ivanova (MHA); Yamalo-Nenetsky 

Autonomous Area: Salemal, Tenguguy 

River, 28.VIII.1963 Soldatenkova (MW). 

NORTH AMERICA: CANADA: Manitoba, S 

of Harashville: Mueller-Dombois #068-13 (MO); New 

Brunswick: Allen #2144 (MO); Newfoundland: 

R.A.C. Waghorne 1.X.[18]93 (MO); Nova Scotia: 

Schofield #97254 (MO); Ontario: O.E. & G.K. Jennings 

s.n. (MO); Prince Edwards Island: Best, Canadian 

Musci, # 380 (MO); Quebeck: Rolland-Germainn 

#115 (MO); U.S.A.: Colorado: Grand Co., 

2750 m (9000’) spruce-fir forest 6 mi W of Fraser, 25 

Sept 1963 Weber et al. (COLO, B-11091), Connecticut: 

26.X.1979 J.A. Allen s.n. (MO), Maine: Allen 

#14770 (MO); Maryland: Austin herbarium 359 coll. 

J.D.Smith (NY); Michigan: Cronin 76065 (NY); 

Montana, Rocky mountains, Columbia Falls: R.S. 

Williams, North American Mosses, #259, 19.XI.1896 

(NY - two specimes with identical label); New Hampshire: 

11.VIII.1878 J.A.Allen (MO); New Jersey: 

Austin Musci Appalaciani, #323 (NY); Nov 1866 

Austun, (NY); New York: Allen #27178 (MO); Pennsylvania: 

Moul 4276 (NY); Vermont: A.J.Grout 

North American Musci Pleurocarpi, 62 (as Brachythecium 

starkei); West Virginia: 1.Sept.1922 Bray 

(NY). 


Sciuro-hypnum oedipodium 

55 

EUROPE: RUSSIA: Caucasus: Karachaevo- 

Cherkessia: Teberdinskij Reserve, Ignatov & Ignatova 

#05-3850, 05-3038 (MHA). 

ASIA: RUSSIA: Chukotka: Gil’mimliveem Hot 

Springs, 20.VII.1977 A.E.Katenin (LE). 

NORTH AMERICA: CANADA: British Columbia: 

Schofield #109936 (MO); Yukon: Willians, Noth 

American Mosses #718 (NY); U.S.A.: Alaska: Darigo 

#2641 (MO); Arizona: Oct 25, 1924 Bartram (NY); 

Colorado: RM King & RMGarvey #B266 (MO); Idaho: 

Plants of Idaho 1297, coll. Sandberg (MO); Montana: 

20.X.1896 R.S. Williams #346; New Mexico: 

Buck #39713 (MO); Utah: Flowers #2635, 2749, 4785; 

Washington: Schofield & Dube #22352 (MO); Wyoming: 

Darigo & Darigo #3275 (MO). 

MOLECULAR DATA 

The protocol of DNA extraction, PCR and sequencing 

of ITS data was the same as described 

in Gardiner et al. (2005). Sequences were aligned 

manually. 

Alignment is presented in the Fig. 7. 

Results can be commented on as follow: 

1. Sciuro-hypnum reflexum, S. starkei, and S. 

oedipodium are very similar in ITS1-2 region, and 

the former species consistently differ from the two 

latter species in just one substitution in ITS2 (specific 

for each of two species). 

2. Sciuro-hypnum curtum is different from S. 

reflexum in 8 substitutions (3 in ITS1 and 5 in 

ITS2), and from S. oedipodium and S. starkei in 9 

substitutions. Available ITS2 data of Huttunen on 

S. curtum from Finland, used in the analysis of 

Huttunen & Ignatov (2004) and not shown in Fig. 

7 has the same 5 characteristic substitutions of 

this species. 

3. Five studied specimens of S. curtum from Europe, 

East Asia and eastern North America are almost 

identical, differing in point mutations in individual 

species, but never corresponding to other 

specimens of this species, or to other species. 

4. There is only very low variation in the group 

of species that includes S. oedipodium, S. reflexum, 

and S. starkei (and also S. altaicum, S. latifolium, 

S. glaciale – see Ignatov & Milyutina, 2007). 

Only one substitution differentiates S. reflexum 

(some populations) and S. starkei, and similarly, 

only one substitution differentiates S. reflexum 

(some populations) and S. oedipodium.


Fig. 7 (see also next page). Alignment of ITS1 &2 and gene 5.8S RNA of Sciuro-hypnum species, demonstrating 

sharp differentiation of S. curtum (8 substitutions, marked by squares) from S. oedipodium, S. reflexum 

and S. starkei, that differs from each other by just one substitution (circles). Specimen data are in Table 1. 

5 15 25 35 45 55 65 75 85 95 105 115 

curtum Tatarstan CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCTCCCCTCG TCACGAGCGC GAGCTTCGTG 

curtum Maine CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCTCCCCTCG TCACGAGCGC GAGCTTCGTG 

curtum New York CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCTCCCCTCG TCACGAGCGC GAGCTTCGTG 

curtum Kamchatka CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCTCCCCTCG TCACGAGCGC GAGCTTCGTG 

curtum Vologda CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCTCCCCTCG TCACGAGCGC GAGCTTCGTG 

reflexum Khabarovsk CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

reflexum Kamchatka CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

reflexum Altai CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

reflexum Vologda CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

oedipodium Canada CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

oedipodium Caucasus CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

oedipodium Alaska CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

oedipodium Wyoming CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

starkei Buryatia CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

starkei Tatarstan CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

starkei Caucasus CACACACAAG TTGCAGCCAA CCCGCGGCGA ACGAACACAT TGTCCCCCCT TCGGCGGGTC GTGGCGCCCT CGCGGCGTCG CGGCCGGCCG TCACCCCTCG TCACGAGCGC GAGCTTCGTG 

125 135 145 155 165 175 185 195 205 215 225 235 

curtum Tatarstan GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTG CCCTCGAGCG AGCTTGCGAA CTCGAGGCGC GCCGCGCGCG 

curtum Maine GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTG CCCTCGAGCG AGCTTGCGAA CTCGAGGCGC GCCGCGCGCG 

curtum New York GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTG CCCTCGAGCG AGCTTGCGAA CTCGAGGCGC GCCGCGCGCG 

curtum Kamchatka GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTG CCCTCGAGCG AGCTTGCGAA CTCGAGGCGC GCCGCGCGCG 

curtum Vologda GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTG CCCTCGAGCG AGCTTGCGAA CTCGAGGCGC GCCGCGCGCG 

reflexum Khabarovsk GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AgcTTGCGAA CTCGAGGCGT GCGGCGCGCG 

reflexum Kamchatka GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

reflexum Altai GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

reflexum Vologda GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

oedipodium Canada GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTSGGTGC GTGTTCCGTS CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

oedipodium Caucasus GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTCGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

oedipodium Alaska GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

oedipodium Wyoming GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

starkei Buryatia GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

starkei Tatarstan GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

starkei Caucasus GCGTCGGGTT TCCACACGAT GCATCACCCG AATGACTGAG TCCCGAATAC TTGGATGGAG GAGTGGGTGC GTGTTCCGTC CCCTCGAGCG AGCTTGCGAA CTCGAGGCGT GCGGCGCGCG 

245 255 265 275 285 295 305 315 325 335 345 355 

curtum Tatarstan CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

curtum Maine CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

curtum New York CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

curtum Kamchatka CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

curtum Vologda CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

reflexum Khabarovsk CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

reflexum Kamchatka CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

reflexum Altai CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAACCA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

reflexum Vologda CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

oedipodium Canada CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

oedipodium Caucasus CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

oedipodium Alaska CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

oedipodium Wyoming CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

starkei Buryatia CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

starkei Tatarstan CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT 

starkei Caucasus CCGGTTACCT TACGACGAGT TGAAACATAA CCAAAAA-CA ACTCTCAGCA ACGGATATCT TGGCTCTTGC AACGATGAAG AACGCAGCGA AATGCGATAC GTAGTGTGAA TTGCAGAATT

Fig. 7. continued from the previous page. 

365 375 385 395 405 415 425 435 445 455 465 475 

curtum Tatarstan CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

curtum Maine CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

curtum New York CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

curtum Kamchatka CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

curtum Vologda CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

reflexum Khabarovsk CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

reflexum Kamchatka CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

reflexum Altai CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

reflexum Vologda CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

oedipodium Canada CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

oedipodium Caucasus CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

oedipodium Alaska CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

oedipodium Wyoming CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

starkei Buryatia CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

starkei Tatarstan CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 

starkei Caucasus CCGCGAATCA TCGAGTCTTT GAACGCAAGT TGCGCCCGAG GCTTGTCCGA GGGCATTTCC GCTAGAGCGT CACCGCGCCC CCCACCGATC CACTCTCGCC GCGAGGCGCC AGTCGGGTCG 


485 495 505 515 525 535 545 555 565 575 585 595 

curtum Tatarstan AGTGCAAGTG GCCGTCCCGG TGGAGCGGCC TCCTTTCACG AGGAGGCCGT ACTCAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

curtum Maine AGTGCAAGTG GCCGTCCCGG TGGAGCGGCC TCCTTTCACG AGGAGGCCGT ACTCAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

curtum New York AGTGCAAGTG GCCGTCCCGG TGGAGCGGCC TCCTTTCACG AGGAGGCCGT ACTCAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

curtum Kamchatka AGTGCAAGTG GCCGTCCCGG TGGAGCGGCC TCCTTTCACG AGGAGGCCGT ACTCAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

curtum Vologda AGTGCAAGTG GCCGTCCCGG TGGAGCGGCC TCCTTTCACG AGGAGGCCGT ACTCAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

reflexum Khabarovsk AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTCACG AGGAGGCCAT ACTGAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

reflexum Kamchatka AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTCACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

reflexum Altai AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTCACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

reflexum Vologda AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTCACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

oedipodium Canada AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTYACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

oedipodium Caucasus AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTTACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

oedipodium Alaska AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTTACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

oedipodium Wyoming AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TCCTTTTACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

starkei Buryatia AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TTCTTTCACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

starkei Tatarstan AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TTCTTTCACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

starkei Caucasus AGTGCAAGTG GCCGTCCCGG TGAAGCGGCC TTCTTTCACG AGGAGGCCAT ACTTAGAAGG GTCGGCTGAA ATGGATGGAA TCGGGCCGCC GTGGTGACAC AGTCCGCGAT CGGGTGATTT 

605 615 625 635 645 655 665 675 685 695 705 715 

curtum Tatarstan TGTCTCGAGC CTCCTTCGGG AGGTTCTTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

curtum Maine TGTCTCAAGC CTCCTTCGGG AGGTTCTTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

curtum New York TGTCTCGAGC CTCCTTCGGG AGGTTCTTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

curtum Kamchatka TGTCTCGAGC CTCCTTCGGG AGGTTCTTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

curtum Vologda TGTCTCGAGC CTCCTTCGGG AGGTTCTTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

reflexum Khabarovsk TGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

reflexum Kamchatka TGTCGCGAGC CTCCTGTGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

reflexum Altai TGTCGCGAGC CTCTTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

reflexum Vologda TGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

oedipodium Canada TGTCGCGAGY CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

oedipodium Caucasus TGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

oedipodium Alaska TGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCACGGCGCG CGTCACGCAT 

oedipodium Wyoming TGTCGCGAGC CYCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

starkei Buryatia CGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGnnnn nnnnnnnnnn 

starkei Tatarstan TGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

starkei Caucasus TGTCGCGAGC CTCCTGCGGG AGGTTCGTGG TGAATTTTTC TAAGTGCTTG GCTGCGTCCC CCAGTTGGCT CGGGTTACCT CATGCAGGTC GAGCGCGCCC CCGCGGCGCG CGTCACGCAT 

57


2 mm 

1 mm 

8 

1 

9 

2 

10 

Fig. 8. Sciuro-hypnum starkei (Brid.) Ignatov & 

Huttunen (from Canada, Ireland 24235, MO): 1 – habit; 

2-3 – mid-leaf cells; 4 – upper leaf cells; 5-7 – branch 

leaves; 8-10 – stem leaves; 11 – basal leaf cells. Scale 

bars: 2 mm for 1; 1 mm for 5-10; 100 μm for 2-4, 11. 

Although the difference is small and includes 

the only substitution between S. reflexum and S. 

starkei, and between S. reflexum and S. oedipodium, 

these differences are consistent with species 

morphology and constant in the territory of the ca. 4 

000 km for S. starkei, and all around Holarctic for S. 

oedipodium. 

WHAT IS SCIURO-HYPNUM STARKEI? 

As most of collections of S. oedipodium and 

S. curtum are still deposited in herbaria (espe- 

5 

6 

3 

100 μm 

11 

cially in North America) under ‘Brachythecium 

starkei’, and this species is considered to be a 

common one by, e.g. Lawton (1971), a brief comment 

on S. starkei is necessary (for standard description 

see Ignatov & Milyutina, 2007). 

Sciuro-hypnum starkei is a medium-sized 

moss that has relatively regular pinnate branching, 

with rigidly spreading branch leaves and 

sometimes also stem leaves. Stem leaves are 

lanceolate-triangular to ovate-triangular, grad- 

7 

4

A 

B 

C 


? 

Fig. 9. Distribution of Sciuro-hypnum curtum (A), S. oedipodium (B) and S. starkei (C), basing on revised 

specimens. Mapping is somewhat schematical, with just one dot per country, except Russia, U.S.A. and Canada, 

where mapping was up to level of state/province or else administrative units. 

ually tapered to apex or sometimes shortly 

acuminate above, narrowly decurrent. Costa is 

vanishing in the narrow part of acumen or often 

percurrent, occasionally ending shortly below 

acumen. Cells at mid-leaf are 40-60 μm, 

but usually with lengh to width ratio ca. 10:1, 

so looking very narrow and somewhat flexuose; 

cells in upper part of branch leaves (occasionally 

in stem leaves also) are prorate on the 

59 

dorsal side. Alar cells are enlarged and thinwalled, 

forming a pellucid group just proximal 

to the decurrency. Seta is always rough; capsules 

are relatively short. Under-developed 

plants differ in more remote and more narrow 

leaves, retaining however their rigidly spreading 

appearance. 

Rigidly spreading leaves and prorate laminal 

cells of S. starkei result in an apprearance quite


similar to Kindbergia praelonga; the latter is different 

however in broader decurrencies, laminal 

cell proportions 4-8:1 (vs. 8-15:1 in S. starkei), 

dioicous sexual condition and longly rostrate operculum. 

SELECTED SPECIMENS EXAMINED: 

Sciuro-hypnum starkei 

EUROPE: RUSSIA: Arkhangelsk Prov.: Onega 

Distr., Ponga, 4.VII.2000 Churakova #80 (MHA); 

Bashkiria: Malyj Iremel Mt., Ignatova #2/111 (MHA); 

Chelyabinsk Prov., Miass Distr., Novoandreevskaya, 

26.VII. 1995 Muldashev #109 (MW); Chuvashskaya 

Republic: Prisursky Nature reserve, 12.VIII.2000 Sultanova 

(MW); Karelia: Olonetsky Distr., Obninsky 

Forestry Unit, 12.VIII.1984 Volkova (LE); Kirov Prov.: 

Khalturin Distr., between Aksenovka and Bakulikha 

(26 km from Nurashi Station) 23.IX.1928 Kolokolnikov 

(LE); Komi Republic: Pechoro-Ilych Nature Reserve, 

27.VI.2000 Bezgodov (Herb. Perm Ped. Univ.); 

Kostroma Prov.: Manturovo Distr., Khalbuzh 

6.VII.1997 Lazareva (MHA); Moscow Prov.: Khimki 

Distr., Yurlovo 5.VIII.1986 Ignatov (MHA); Murmansk 

Prov., Polar-Alpine Botanical Garden, 500 m, 

31.VII.1998 Ignatov & Ignatova (MHA); Nizhniy 

Novgorod Prov.: Kerzhensky Reserve, 100 m, 

18.IX.1999 Ignatov (MHA); Novgorod Prov.: Valdai 

Distr., 25.VII.1981 Vaulina (MHA); Perm Prov.: 

Vishera State Reserve, 20.VII.1994 Bezgodov (MHA); 

Sverdlovsk Prov.: Kirovgrad Distr., Visimsky Nature 

Reserve, 9.VII.2005 Smirnova (MW); Tatarstan, 

Volzhsko-Kamsky Reserve, Saraly, Ignatov & Ignatova 

#05-2131 (MHA); Tver Prov.: Centralno-Leskoj Reserve, 

200 m, 28.VIII.1989 Minaeva (MHA); Udmurtia: 

Votkinsk Distr., Kostovaty 17.VII.2000 Munitsyna 

(MHA); Vladimir Prov: Aleksandrovo Distr., Seregin 

& Privalova #M-388 (MW); Vologda Prov.: Sokol 

Distr., Shorega Creek, 22.IX.1990 Ignatov (MHA); 

Yaroslavl Prov.: Pleshcheevo Lake, 20.VIII.2001 Ignatov 

& Ignatova (MHA). CAUCASUS: Karachaevo-Cherkessia: 

Teberdinsky Reserve, 2850 m, 

8.IX.2005 Ignatov & Ignatova (MHA); Krasnodar 

[AFONINA, O.M.] АФОНИНА О.М. 2004a. Конспект 

флоры мхов Чукотки. – [Moss flora of Chukotka] СПб, 

БИН РАН [Sankt-Petersburg, Bot. Inst. RAS], 260 pp. 

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of mosses of North America north of Mexico. – Bryologist 

93: 448-499. 

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ITSKY 2005. On resurrection of the families Pseudoleskeaceae 

Schimp. and Pylaisiaceae Schimp. (Musci, Hyp- 

LITERATURE CITED 

Territory: Caucasian Nature Reserve, Abago Mt., 1850 

m, 20.VII.1960 Artamonova (MW). GEORGIA: South 

Osetia, Dzava Distr., 13.VIII.1955 Dombrovskaya 

(MW). GERMANY: Bayrisches Wald, 20.X.1918 Paul 

(MW). POLAND: Zarnowiec & al.: Musci Macroregioni 

Merid. Poloniae Exs. #302 (MHA). 

ASIA: RUSSIA: Altai Territory: Riddera, 

25.VIII.1947 Polyakov (MHA); Altai Republic: Kairu-Bazhi 

Peak, 2050 m, Ignatov #13/105 (MHA); 

Amurskaya Territory: Zeya Nature Reserve, 15.VIII.1980 

L.Abramova #126 (MW); Buyatia: Khamar-Daban, 

Vydrinnaya, 15.VIII.1989 Kazanovsky (MHA ex 

IRK); Evenkiya: 62°09’N,91°09’E, 19.VIII.1992 Szerbina 

(MW); Irkutsk Prov.: Ulkan, 1.VII.1976 Bardunov 

(MHA ex IRK); Kamchatskaya Prov.: 

Kozyrevsk, 220 m, 12.VIII.2004 Czernyadjeva #64 

(LE); Kemerovo Prov.: Tisulsky Distr., Kiya River, 400 

m, 7.IX.1971 Vasiliev (MHA ex IRK); Khabarovsk 

Territory: De Castri Bay, 18.VI.1854 Maximovicz (LE); 

Primorsky Territory: Lazo Distr., Olkhovaya Mt., 1650 

m, 2.X.2006 Ignatov, Ignatova & Cherdantseva #06- 

2200 (MHA); Sakhalinskaya Prov.: Sakhalin, Dui, V- 

1861 Glehn (LE); Kunashir, Saratovka, Ignatov #06- 

1387 (MHA). 

NORTH AMERICA: CANADA: New Brunswick: 

Ireland 13008 (MO); Labrador: R.A.C.Waghorne 

1.X.[18]93 (MO); Ontario: Ireland 24235 (MO); Prince 

Edwards Island: J. Macoun Canadian Mosses # 293 

(MO); Quebeck: Redfearn 37440 (MO); U.S.A.: Maine: 

Allen 14774, 24555 (MO); Vermont: E.C.Taylor S-697 

(NY); Wisconsin: Pursell 6120 (MO). 

ACKNOWLEDGEMENTS 

We thank curators of H, IRK, LE, MW, NVS, 

S, SASY, VLA for making their respective collections 

available for study; Richard Zander and 

William Buck for arranging the visit of the first 

author to MO and NY. We also thank Elena Ignatova 

for preparing illustrations of species and 

Angela E. Newon for the English correction. The 

work was partly supported by the RFBR, grants 

07-04-00013 and 06-04-49493. 

nales). – Taxon 54(3): 651–663. 

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GROUT, A. J. 1928. Moss flora of North America, north of 

Mexico. Vol. 3, Pt. 1, Published by the author, Newfane, 

Vt. & New-York: 1-62.


Table 1. Specimen data of the genus Sciuro-hypnum used in analysis, and Genbank accession numbers. 

species specimen GenBank # 

curtum Kamchatka Asian Russia, Kamchatka, Czernyadjeva 48-07 (MHA) EU715332 

curtum Maine U.S.A., Maine, Allen 14770 (MO) EU715330 

curtum New York U.S.A., New York State, Allen 27178 (MO) EU715331 

curtum Tatarstan European Russia, Tatarstan, Ignatov & Ignatova 18.VIII.2003 (MHA) EU715329 

curtum Vologda European Russia, Vologda, Ignatov & Ignatova 19-8-2001 (MHA) EU715333 

oedipodium Caucasus Russia, Caucasus, Ignatov & Ignatova 05-3038 (MHA) EU715345 

oedipodium Alaska U.S.A., Alaska, Darigo 2641 (MO) EU715346 

oedipodium Wyoming U.S.A., Wyoming, Darigo 3275 (MO) EU715347 

oedipodium British Columbia Canada, British Columbia, Schofield 109936 (MO) EU715344 

reflexum Kamchatka 39 Asian Russia, Kamchatka, Chernyadjeva 5.VIII.2002 #39 (MHA) EU715351 

reflexum Khabarovsk Asian Russia, Khabarovsk Territory, Ignatov 97-177 (MHA) EU715352 

reflexum Vologda European Russia, Vologda Province, Ignatov & Ignatova 19.VIII.2001 (MHA) EU715353 

reflexum ‘pacificum’ Altai Asian Russia, Altai, Ignatov 0/442 (MHA) EU715354 

starkei Tatarstan European Russia, Tatarstan, Ignatov & Ignatova 05-2027 (MHA) EU715348 

starkei Caucasus Russia, Caucasus, Onipchenko 31-8-1983 #37-83 (MHA) EU715350 

starkei Buryatiya Asian Russia, Buryatia, Kazanovsky 16.VIII.1989 (MHA) EU715349 

GROUT, A. J. 1934. Moss flora of North America, north of 

Mexico. Vol. 3, Pt. 4, Published by the author, Newfane, 

Vt. & New-York: 179-277. 

HILL, M.O., N. BELL, M.A. BRUGGEMAN-NANNENGA, 

M. BRUGUÉS, M.J. CANO, J. ENROTH, K.I. FLAT- 

BERG, J.-P. FRAHM, M.T. GALLEGO, R. GARILLETI, 

J. GUERRA, L. HEDENAS, D.T. HOLYOAK, J. HYVÖ- 

NEN, M.S. IGNATOV, F. LARA, V. MAZIMPAKA, J. 

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of the mosses of Europe and Macaronesia. – J. Bryol. 

28: 198–267. 

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of Brachytheciaceae (Bryophyta) based on morphology, 

and sequence level data. – Cladistics 20: 151-183. 

IGNATOV, M. S. 1998. Bryophyte flora of Altai Mountains. 

VIII. Brachytheciaceae. — Arctoa 7: 85-152. 

IGNATOV, M.S., O.M.AFONINA, E.A. IGNATOVA et al. 

2006. Check-list of mosses of East Europe and North Asia. 

– Arctoa 15: 1-130. 

61 

IGNATOV, M.S. & S. HUTTUNEN 2002. Brachytheciaceae 

(Bryophyta) – a family of sibling genera. – Arctoa 11: 245- 

296. 

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the genus Sciuro-hypnum (Brachytheciaceae, Bryophyta) 

in Russia. – Arctoa 16: 73-94. 

LAWTON, E. 1971. Moss Flora of the Pacific Northwest. – 

Nichinan, Hattori Bot. Lab., xiii + 362 pp. + 195 pl. 

LIMPRICHT, K. G. 1896. Die Laubmoose. – In: Rabenhorst, 

L. Kryptogamen-flora von Duetschland, Oesterreich und 

der Scheiz. Bd. 3. 1-448. 

PIIPPO, S. 1983. On the taxonomy, nomenclature and distribution 

of Brachythecium starkei (Brachytheciaceae, 

Musci) and related taxa. — Ann. Bot. Fennici 20: 339- 

349. 

REDFEARN, P.L., JR., B.C.TAN & S. HE 1996. A newly 

updated and annotated checklist of Chinese mosses – J. 

Hattori Bot. Lab. 79: 163-357.

ON SCIURO-HYPNUM OEDIPODIUM AND S. CURTUM - Arctoa

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