Key to European Dicranum species

Abstract

HEDENÄS, L. & BISANG, I. 2004. Key to European Dicranum species. - Herzogia 17: 179-197. A key to the twenty-nine European Dicranum species is presented. The features that are important for their identifi- cation are discussed. For each species, diagnostic character states and information about habitat and geographical distribution are provided. Most critical characters are illustrated. Zusammenfassung: HEDENÄS, L. & BISANG, I. 2004. Bestimmungsschlüssel für die Dicranum-Arten Europas. - Herzogia 17: 179-197. Ein Schlüssel für die neunundzwanzig Dicranum-Arten Europas wird vorgestellt. Die zur Bestimmung der Arten wich- tigen Merkmale werden diskutiert. Für jede Art werden die diagnostischen Merkmale beschrieben sowie Standortsan- sprüche und geographische Verbreitung dargestellt. Die meisten der kritischen Merkmale sind abgebildet.

Full text

HEDENÄS & BISANG: Key to European Dicranum species 179
Herzogia 17 (2004): 179–197
Key to European Dicranum species
Lars HEDENÄS & Irene BISANG
Abstract: HEDENÄS, L. & BISANG, I. 2004. Key to European Dicranum species. – Herzogia 17: 179–197.
A key to the twenty-nine European Dicranum species is presented. The features that are important for their identifi-
cation are discussed. For each species, diagnostic character states and information about habitat and geographical
distribution are provided. Most critical characters are illustrated.
Zusammenfassung: HEDENÄS, L. & BISANG, I. 2004. Bestimmungsschlüssel für die Dicranum-Arten Europas. –
Herzogia 17: 179–197.
Ein Schlüssel für die neunundzwanzig Dicranum-Arten Europas wird vorgestellt. Die zur Bestimmung der Arten wich-
tigen Merkmale werden diskutiert. Für jede Art werden die diagnostischen Merkmale beschrieben sowie Standortsan-
sprüche und geographische Verbreitung dargestellt. Die meisten der kritischen Merkmale sind abgebildet.
Key words: Dicranaceae, identification guide, mosses.
Introduction
Members of the genus Dicranum are widespread all over Europe. At the same time some species
have become rare, and in some cases even endangered in those parts of the continent that are
most heavily influenced by human activities. Most of the almost thirty European species are
relatively large in size and several of them frequently cover large patches of their substrates.
Many species are therefore quantitatively important, especially in forest and mountain habitats.
A number of species seem to have special and relatively narrow habitat requirements and can
therefore function as indicators for specific conditions, at least in certain regions.
In view of this, the role of Dicranum species in nature appears to be an evident target for in-
depth studies in the fields of ecology and nature conservation. However, the frequently con-
tradicting information that is provided in the literature regarding which species to recognise,
how to separate several species, and the frequently quantitative distinguishing characters of-
ten hamper the identification of members of Dicranum.
In the present overview, a total of twenty-nine species are recognised in Europe. Problematic
taxa are discussed in the notes after the key.
Important characters
Species of Dicranum are usually recognised as such by their typically large size and their nar-
row, lanceolate leaves that are more or less gradually narrowed towards the leaf apex, a long,
single costa, and in most species well differentiated alar groups. The leaves may be straight, but
are frequently homomallous to falcate, and the colour is mostly a rather deep and clear green.
Members of a few other genera of the family Dicranaceae are also relatively large, and are
superficially similar to Dicranum species in habit. Most species of Dicranodontium, Campylopus
and Paraleucobryum have got broader costae than most Dicranum species, and in addition the

180 Herzogia 17 (2004)
anatomy of the costa is different (NYHOLM 1987). A northern European species that is now and
then misidentified as a Dicranum is Kiaeria glacialis (Berggr.) I. Hag. (fig. 1a), but this species
is autoicous and has differently organised alar cells (fig. 1b) compared to those Dicranum spe-
cies with which it could be confused. Especially, the alar cells are unistratose (in Dicranum
found only in D. montanum, D. flagellare, and D. tauricum), and the inner alar cells never get
thin and translucent (fig. 1c) like in the Dicranum species with which it can be confused.
Fig. 1: a – Kiaeria glacialis, habit (moist), b – alar groups, c – Dicranum angustum, alar groups.
Capsule orientation
Within the genus Dicranum, few sporophytic characters are useful for species identification. Since
Dicranum species are dioicous and many species therefore relatively rarely produce sporophytes,
one cannot rely only on sporophytic features for their identification. Thus, a key that separates
1
/
5
of the species from the rest only by the orientation of the spore capsule (fig. 2), such as the one in
N
YHOLM (1987), is useless for most specimens.
Dwarf male plants
Many Dicranum species have dwarf male plants (fig. 3) that grow on or
among the leaves, or in the frequently present dense rhizoids on the stems of
the female plants. Male plants of about the same size as the female plants
are found in D. elongatum, D. flagellare, D. flexicaule, D. fragilifolium, D.
fulvum, D. fuscescens, D. groenlandicum, D. montanum, D. scottianum, D.
tauricum, and D. viride, and probably in D. canariense and D. laevidens as
well. In D. scoparium, both dwarf and normal-sized males occur, while the
remaining species have only dwarf male plan
ts.
Plant size
A feature that aids in the identification of some species is the overall
plant size. While among most species the plants are of approximately
the same size, those of D. majus (fig. 43), D. drummondii and D.
polysetum are generally markedly larger than in the other species, whereas
D. montanum (fig. 44) and D. flagellare have considerably smaller plants
than the remaining ones.
Flagellae
Flagellae are shoots or shoot portions with small and more or less closely
imbricate leaves. Such shoots may arise in the leaf axils or may consist
of the upper, transformed parts of shoots that look normal below.
Flagellae occur with different frequencies in all Dicranum species. In
species such as D. leioneuron and D. flagellare they are almost invari-
ably present, in other ones, like D. bonjeanii and D. scoparium, they
are seen now and then, while in most species they are rare.
a b c
Fig. 3: Dwarf male plant in
Dicranum spurium.
Fig. 2: a – Straight and erect
spore capsule in Dicranum
scottianum; b – curved,
asymmetric capsule in D.
scoparium.
a b

HEDENÄS & BISANG: Key to European Dicranum species 181
Vegetative leaf characters
Most of the characters that are useful for the identification of Dicranum taxa are found in the
vegetative leaves. Besides leaf shape and whether the lamina is transversely undulate or not,
a number of features related to the lamina cells are important. We would like to stress that it is
important to study numerous leaves in order to correctly judge their characters. This espe-
cially concerns the lamina cell appearance.
Leaf orientation. Leaf orientation is one of the most useful characters for the recognition of
many species, especially in the field. When moist, species like D. undulatum have mostly got
all leaves erect, whereas in D. polysetum only the upper leaves are erect, and most leaves are
spreading. Both these species have got straight or slightly curved leaves whereas, for exam-
ple, D. majus and D. fuscescens have mostly got strongly falcate leaves. In some species, such
as D. scoparium, straight- and falcate-leaved types are both frequent. Leaf orientation fre-
quently differs between moist and dry plants, for example, in D. brevifolium, which has more
or less secund leaves when moist and strongly curled ones when dry.
Transverse section of leaf and costa. The shape of transverse sections made in the upper
portion of the leaves is sometimes diagnostic. In some cases, such as when tubular and flat
leaves are compared, the shape of the section can be guessed from observations of the entire
leaves. In other cases, it is necessary to really make sections, for example, to see the tong-
shaped sections of D. brevifolium and a few other species. Also the costal anatomy (fig. 4) can
be crucial for the recognition of certain species. Especially, the presence or absence of stereid
cells and how well the epidermal cells are differentiated is of interest. In a few species two or
more lamellae are present on the back of the costa. Such structures are well developed in D.
scoparium and closely related taxa.
Leaf lamina cells. Starting from the leaf base, the alar cells are usually at least partly bistratose,
but as mentioned above, D. montanum, D. flagellare, and D. tauricum have unistratose, and
D. dispersum has partly 3(–4)-stratose alar groups. Also the lamina cells further up may be
bistratose, and are regularly so in several species, at least along the upper margins. Cell size is
usually either too variable or too similar among species that are problematic to separate to be
useful for their identification. On the other hand, both cell shape and cell wall thickness are
frequently diagnostic. Both these features are sometimes difficult to estimate without some
experience, and have therefore been illustrated in the key when relevant. Other lamina cell
features that are sometimes important for the recognition of species are the frequency and
development of pores in the cell walls, whether mammillae or spines are present on the dorsal
side of the lamina, and whether the cell walls are projecting (seen in transverse sections of the
lamina).
Leaf margin. In the present key denticulate means that cell portions form the ‘teeth’ of the
leaf margin, whereas dentate implies that entire cells form the ‘teeth’.
Fig. 4: Transverse section of costa in a – Dicranum tauricum, b – D. viride; c – D. scottianum.
a b c

182 Herzogia 17 (2004)
Key to the European Dicranum species
1 Leaves stiff and fragile, in at least upper part of shoot straight
and erect (
fig. 5a), upper leaf portions very narrow and tips fre-
quently broken (
fig. 5b) ........................................................ 2
1* Leaves in upper part not at the same time fragile, straight and
erect, upper leaf portions not markedly narrow ..................... 4
2 Costa in basal portion lacking stereid bands (
fig. 4a), with up to
1–2 cell layers above and below guide cells; basal lamina cells
narrowly rectangular to linear, eporose; alar cells unistratose
(
fig. 6). Capsule straight and erect, or almost so ...... D. tauricum
2* Costa in basal portion with stereid bands, sometimes weak, with
up to 2–3(–4) layers of cells above and below guide cells; basal
lamina cells elongate and porose or rectangular and eporose;
alar cells mostly bistratose at least in parts, rarely entirely
unistratose. Capsule straight and erect or asymmetrical and
curved ..................................................................................... 3
3 Epidermal cells of costa not differentiated from stereids; basal
lamina cells narrowly rectangular to linear, more or less porose;
lamina mostly unistratose. Capsule asymmetrical and more or
less curved ......................................................... D. fragilifolium
3* Epidermal cells of costa wider than stereids, at least on dorsal
side; basal lamina cells rectangular, not or hardly porose; lamina
above in parts bistratose. Capsule straight and erect, or almost
so ................................................................................... D. viride
4 Cells in upper leaf mainly prosenchymatous, elongate and porose
(
fig. 7) ................................................................................... 5
4* Cells in upper leaf mainly parenchymatous, quadrate or rectan-
gular, rarely elongate-rectangular, with or without pores (fig.
8). (Note that in D. spadiceum the parenchymatous cells are
sometimes restricted to the uppermost part of the acumen, at
least in some leaves) ............................................................. 13
5 Leaves straight, in upper part more or less tubular (
fig. 9a; trans-
verse section); apex rounded (
fig. 9b) or obtuse, sometimes acute
(
fig. 9c); margin entire throughout or sometimes with a few ob-
tuse denticles near apex ........................................................ 6
5* Leaves falcate or straight, in upper part flat or channeled (e.g.
fig. 12), apex acuminate; margin above finely to coarsely den-
ticulate .................................................................................... 8
6 Plants in dense, compact tufts (usually so dense that it is diffi-
cult to press your finger into the tuft). Peristome papillose
........................................................................ D. groenlandicum
6* Plants in loose tufts. Peristome papillose or smooth .............. 7
7 Almost invariably with flagellae having small, crowded, imbri-
cate leaves. Leaf acumen relatively short and broad; costa with
or without two weak and, when seen in transverse section, ob-
tuse ridges. Peristome papillose ........................... D. leioneuron
7* Flagellae extremely rare. Leaf acumen long and narrow; costa
without ridges. Peristome smooth ........................... D. laevidens
Fig. 5a
(dry)
Fig. 5b
(dry)
Fig. 6
Fig. 7
Fig. 8

HEDENÄS & BISANG: Key to European Dicranum species 183
8 Costa above with two lamellae or ridges on back .................. 9
8* Costa above with at least four lamellae or numerous ridges or
furrows on back (at least in some leaves) ............................ 10
9 Costal lamellae tall; leaf lamina strongly transversely undulate;
margin in upper part spinosely denticulate or dentate. Apical
leaves erect, leaves below spreading (
fig. 10) ....... D. polysetum
9* Costal lamellae low or weakly developed; leaf lamina more or
less strongly transversely undulate (at least in some leaves); mar-
gin obtusely to sharply denticulate. Orientation of apical leaves
and leaves further down not clearly different .......... D. bonjeanii
10 Costa above with numerous furrows or low ridges; leaf lamina
cells in transverse section mostly rectangular (longest in direc-
tion from costa to leaf margin) ...................................... D. majus
10* Costa above with distinct lamellae (
fig. 11; right: upper costa
seen from dorsal side); leaf lamina cells in transverse section
mostly approximately quadrate ............................................. 11
11 Leaf lamina unistratose ......................................... D. scoparium
11* Leaf lamina above partly bistratose, or in spots tristratose ............. 12
12 Leaf lamina spinose on back (
fig. 12); upper leaf margin spinose-
dentate ......................................................... D. transsylvanicum
12* Leaf lamina smooth (
fig. 13); upper leaf margin denticulate to
dentate ................................................................ D. crassifolium
13 Upper leaf lamina in at least some leaves rugose, or more or
less transversely undulate (dry and wet) .............................. 14
13* Upper leaf lamina neither rugose nor undulate .................... 16
14 Leaf lamina rugose, in upper leaf on back with spine-like (
fig.
14) or conically projecting cells; basal leaf portion ovate, leaf
apex acuminate (
figs 15b, c) ................................................ 15
14* Leaf lamina transversely undulate, in upper part with smooth or
lowly mammillose cells; basal leaf portion lanceolate, leaf apex
more or less obtuse (
fig. 15a) ................................ D. undulatum
15 Upper leaf lamina cells irregularly quadrate, walls esinuose;
leaves gradually narrowed to long, falcate acumen (
figs 15b, 16)
...................................................................... D. drummondii
15* Upper leaf lamina cells very irregular, triangular, quadrate, with
sinuose walls; leaves suddenly narrowed to short acumen (
fig.
15c) (shoot apices when dry and viewed from above reminding
about lettuce heads;
fig. 17) ...................................... D. spurium
Fig. 9
a
b
c
Fig. 10 (moist)
Fig. 11
Fig. 12 (after
LÜTH 2002)
Fig. 13 (after
SÉRGIO et al. 1995) Fig. 14 (left) a b c
Fig. 15:
Stem leaves of
a – Dicranum undulatum,
b – D. drummondii,
c – D. spurium.

184 Herzogia 17 (2004)
16 Small plants (rarely above 3 cm high; leaves up to 4 mm long).
Dry leaves strongly crisped; upper lamina cells mammillose,
normally unistratose (
fig. 18; transverse section of upper lamina)
............................................................................... D. montanum
16* Plant size variable. Leaf orientation variable; upper lamina cells
variable, if mammillose or papillose, then lamina partly
bistratose, leaves fragile, or plants larger ............................. 17
17 Small plants that almost invariably have numerous easily de-
tached flagellae from upper leaf axils (
fig. 19). Alar cells
unistratose .............................................................. D. flagellare
17* Plant size variable; if flagellae present, then never several on
each plant. Alar cells at least partly bistratose ..................... 18
18 Upper leaf lamina bistratose (
fig. 20), sometimes only along mar-
gins (rarely restricted to uppermost margins); leaves frequently
falcate, when dry more or less strongly crisped .................. 19
18* Leaf lamina unistratose (occasional plants may have bistratose
spots); leaf orientation variable ............................................. 26
19 Upper leaf lamina bistratose in large portions (also besides leaf
margins;
fig. 20). Costa very broad (c.
1
/
3
of basal leaf width)
..................................................................................... D. fulvum
19* Leaf lamina bistratose only along margins. Costa narrower, or
in D. scottianum and D. canariense very broad ................... 20
20 Transverse section of upper leaf looking like a pair of tongs (
fig.
21) ....................................................................................... 21
20* Transverse section of upper leaf varying, not looking like a pair
of tongs .................................................................................. 23
21 Plants relatively large (stem leaves 6.0–8.5 mm long). Alar cells
2–3(–4)-stratose (
fig. 22a); epidermal cells on ventral side of
costa differentiated (
fig. 22b); leaf lamina cell walls not project-
ing in transverse section ................................... D. dispersum
21* Plants smaller (stem leaves 3.5–7.0 mm long). Alar cells 2-
stratose; epidermal cells on ventral side of costa undifferenti-
ated or at most slightly differentiated; leaf lamina cell walls pro-
jecting in transverse section (fig. 37) .................................. 22
22 Upper leaf lamina cells quadrate. Upper leaves when dry curled,
leaves when moist usually secund (fig. 23) ......... D. brevifolium
Fig. 16 (dry)
Fig. 17 (dry)
Fig. 18
(moist)
Fig. 19 (moist)
Fig. 21
Fig. 22
a
Fig. 20
b

HEDENÄS & BISANG: Key to European Dicranum species 185
22* Upper leaf lamina cells irregular, quadrate, triangular, or shortly
rectangular. Upper leaves when dry slightly curled to slightly
bent, leaves when moist not or slightly secund .... D. acutifolium
23 Lamina cells thin-walled or slightly incrassate (
fig. 24a). Cap-
sule asymmetrical and more or less curved ......................... 24
23* Lamina cells above incrassate (
fig. 24b). Capsule straight and
erect, or almost so ................................................................. 25
24 Upper leaf margin more or less densely denticulate in upper
1
/
3
–
1
/
2
(fig. 25a), from apex and some distance below frequently
coarsely denticulate; upper lamina cells relatively regularly
quadrate (
fig. 25b), frequently mammillose; costa dorsally
densely denticulate or rough in upper
2
/
5
–
4
/
5
(fig. 25c;
1
/
3
way
down from leaf apex) ............................................ D. fuscescens
24* Upper leaf margin irregularly denticulate, with long entire por-
tions, in upper
1
/
5
–
1
/
3
(fig. 26a), near apex sometimes coarsely
denticulate; upper lamina cells irregular in shape (
figs 26b, c),
only scattered ones sometimes mammillose; costa dorsally rela-
tively distantly denticulate or rough in upper
1
/
3
(fig. 26d;
1
/
3
way
down from leaf apex) ............................ .......... D. flexicaule
25 Leaf margin entire or with few and obtuse denticulations (
fig.
27a) except close to apex, where a few denticles may be present
(
fig. 27b), costa smooth (fig. 27c;
1
/
4
way down from leaf apex),
occasionally with a few denticles near apex; costa near base
(above alar groups) weakly convex on back, with 1–2 ventral
and 2–4 dorsal layers of stereids. Aristate acumen of perichaetial
leaves entire, occasionally weakly denticulate in uppermost part.
Plants when dry slightly glossy ............................ D. scottianum
25* Leaf margin denticulate or dentate in upper (
1
/
7
–)
1
/
6
–
1
/
4
(figs 28a,
b), costa rough in upper
1
/
4
–
1
/
2
(fig. 28c;
1
/
4
way down from leaf
apex); costa near base convex on back, with 2–4 ventral and 3–
5 dorsal layers of stereids. Aristate acumen of perichaetial leaves
obtusely denticulate or denticulate in upper
2
/
5
–
4
/
5
, near apex
partly spinosely denticulate. Plants when dry dull
.......................................................................... D. canariense
Fig. 23 (dry)
a b c d
Fig. 26
Fig. 24 a b
a b c
Fig. 27
a b c
Fig. 28
Fig. 29
Fig. 25 a b c

186 Herzogia 17 (2004)
26 Transverse section of upper leaf looking like a pair of tongs (fig.
29) ......................................................... ........................ 27
26* Transverse section forms a more or less circular outline (costa
plus lamina;
fig. 30a); upper leaf tubular (fig. 30b) .............. 29
27 Plants relatively large (stem leaves 6.0–8.5 mm long). Alar cells
2–3(–4)-stratose (see
fig. 22a); epidermal cells on ventral side
of costa differentiated (see
fig. 22b); leaf lamina cell walls not
projecting in transverse section ............................. D. dispersum
27* Plants smaller (stem leaves 3.5–7.0 mm long). Alar cells 2-
stratose; epidermal cells on ventral side of costa undifferenti-
ated or at most slightly differentiated; leaf lamina cell walls pro-
jecting in transverse section (
fig. 37) .................................. 28
28 Upper leaf lamina cells quadrate. Upper leaves when dry curled,
leaves when moist usually secund (
fig. 23) ......... D. brevifolium
28* Upper leaf lamina cells irregular, quadrate, triangular, or shortly
rectangular. Upper leaves when dry slightly curled to slightly
bent, leaves when moist not or slightly secund .... D. acutifolium
29 Leaves when dry strongly curled. Leaf margin near apex irregu-
larly and coarsely denticulate to dentate ........ D. muehlenbeckii
29* Leaves when dry straight or weakly curled in their upper por-
tions. Leaf margin weakly denticulate to almost entire ........ 30
30 Plants in dense, compact tufts (
fig. 31a). Upper leaf lamina cells
strongly incrassate (
fig. 31b); leaf base lanceolate
................................................................................ D. elongatum
30* Plants more loosely tufted. Upper leaf lamina cells thin-walled
to moderately incrassate (
fig. 32); leaf base ovate or ovate-lan-
ceolate .................................................................................... 31
31 Leaves erect-spreading to spreading (
fig. 33a). Leaf lamina cells
not or hardly porose (
fig. 33b, from basal leaf); costa weak.
Shoots loosely tufted or scattered among other mosses in rich
fens ........................................................................ D. angustum
31* Leaves erect-spreading to loosely appressed (
fig. 34a). Leaf
lamina cells densely porose, at least in lower leaf (
fig. 34b, from
basal leaf); costa strong. Shoots rather closely tufted on moist
soil, rocks, decaying logs, etc. ............................... D. spadiceum
a (moist)
Fig. 32
Fig. 33 a (moist) Fig. 34 a (moist)
b
Fig. 30
a
b
b
Fig. 31
b

HEDENÄS & BISANG: Key to European Dicranum species 187
European Dicranum species
Dicranum acutifolium (Lindb. & Arnell) C. E. O. Jensen [D. bergeri var. acutifolium Lindb.
& Arnell; D. muehlenbeckii var. acutifolium (Lindb. & Arnell) Nyholm]
D. acutifolium is closely related and very similar to D. brevifolium. Both these species, as well
as D. dispersum, are easily recognised by having a tong-shaped transverse section in the upper
leaf portion. The leaves of D. acutifolium are somewhat straighter than those in D. brevifolium,
both when moist and dry, and the upper leaf lamina cells are more irregular than in the latter.
The features distinguishing D. dispersum are mentioned under the latter.
This species is widespread in N and probably also C Europe (numerous finds of D. acutifolium
from Austria in S). – Grows on more or less calcareous soil or rocks, mainly in mountainous areas.
Dicranum angustum Lindb.
This species forms loose tufts in wetlands, or sometimes grows scat-
tered among other bryophytes. It can be differentiated from other
wetland Dicranum species by its long, narrow, tubular, acuminate or
occasionally blunt leaves that are erect-spreading or spreading. The
lamina cells are thin-walled or only slightly incrassate, and are eporose
or only slightly porose, and the leaf margins are entire or only near
apex slightly denticulate (fig. 35).
D. angustum is widespread in the boreal to arctic areas of N Europe. It
is unlikely that it occurs in C Europe. – Grows in more or less mineral-
rich fens.
Dicranum bonjeanii De Not. (D. amannii Peterson, nom. inval.; D. latifolium J. J. Amann; D.
palustre Bruch & Schimp.; D. undulatum Turner)
Among the species having prosenchymatous upper leaf lamina cells, D. bonjeanii can usually
be recognised by its weakly to strongly transversely undulate upper leaf portions (at least in
some leaves), its two weak dorsal leaf lamellae, which may be serrate or entire, and by its leaf
acumen above typically narrowing shortly to a broad, often blunt or obtuse leaf apex. Note,
however, that the length of the acumen may vary from relatively short, common in open or sparsely
tree-covered habitats, to long, usually in swampy forests. Like many other Dicranum species, this
species can have both straight and falcate leaves. In some floras (e.g. NYHOLM 1987) it is stated
that the relatively similar species D. bonjeanii and D. leioneuron can be
separated by the presence of flagellae in D. leioneuron and the absence of
such shoots in D. bonjeanii. However, even if flagellae are almost in-
variably present in D. leioneuron, they occur also in D. bonjeanii (fig.
36) where they seem to be especially common in habitats that dry out
occasionally, such as depressions on calcareous ground. D. leioneuron
is a species of poor fens or bogs and has entire or almost entire leaf
margins, whereas D. bonjeanii occurs in richer habitats and has den-
ticulate or at least obtusely denticulate leaf margins. Also in the ha-
bitually similar D. scoparium, flagellae are occasionally found.
This species is widespread in Europe. – Found in more or less mineral-
rich habitats, such as rich fens or relatively rich mire portions in mixed
mires, moist soil, or in temporarily wet depressions on flat, horizontal
rocks, more rarely in humid non-wetland habitats (SAUER 1999).
Fig. 36: Dicranum
bonjeanii, habit, with
flagellae (moist).
Fig. 35:
Dicranum
angustum, leaf
acumen.

188 Herzogia 17 (2004)
Dicranum brevifolium (Lindb.) Lindb. (D. muehlenbeckii var. brevifolium Lindb.; D.
muehlenbeckii auct., p.p.)
D. brevifolium is usually easily recognised when it is dry, due to its curled or twisted, rela-
tively stiff leaves. Like in D. acutifolium and D. dispersum, transverse sections made in the
upper leaf are tong-shaped in outline. Differences between these three species are discussed
under D. acutifolium and D. dispersum, respectively. AMANN (1918) considered D. brevifolium
a high altitude variant of D. muehlenbeckii, and believed there were transitions between typi-
cal D. muehlenbeckii and D. brevifolium. However, the two are easily distinguished by the
very different appearances of transverse sections made in the upper portions of the leaves,
tong-like in D. brevifolium, circular in D. muehlenbeckii, and
the longitudinal leaf lamina cell walls are strongly projecting in
D. brevifolium (fig. 37), not so in D. muehlenbeckii.
D. brevifolium is widespread in large areas of N, C, and E Eu-
rope. – Grows on usually dry soil or rocks, and is most frequent
in open and calcareous environments.
Dicranum canariense Hampe ex Müll. Hal.
D. canariense belongs to the Dicranum species with erect sporophytes, which in addition are
relatively frequent in this species. It is most closely related to D. scottianum, from which it
differs by much more strongly denticulate leaf margins and costa, and by a thicker costa.
This species occurs on the Canary Islands and Madeira, and is also known from Spain and
possibly France (REIMERS 1930) on the European mainland. – Grows both as an epiphyte, on
Madeira often on Erica scoparia L. and on vertical rocks.
Dicranum crassifolium Sérgio, Ochyra & Séneca
With its partly bistratose or in spots tristratose leaf lamina and distinct lamellae on the upper
back of its costa, this species could only be confused with D. transsylvanicum. The latter has,
however, spinosely dentate rather than denticulate to dentate leaf margins, and spines on the
back of its upper lamina (smooth in D. crassifolium). D. scoparium differs from both these
species in having an unistratose leaf lamina.
SÉRGIO et al. (1995) reported D. crassifolium from Portugal, Spain (up to 1750 m a.s.l.), and
Italy (Toscana, Apuan Alps). – Grows on soil or humus in open or shaded places, as well as on
rocks and at tree bases, preferably in relatively acid places (SÉRGIO et al. 1995).
Dicranum dispersum Engelmark
D. dispersum is a relatively large species, forming tufts that can reach 10 cm height, with straight or
slightly flexuose leaves that are loosely crisped when dry. Like in D. acutifolium and D. brevifolium,
the transverse section is tong-shaped in the upper portion of the leaves. Both the latter species can be
separated from D. dispersum by their not or poorly differentiated ventral epidermal costa cells, by
thickened longitudinal cell walls (fig. 37), and by never having 3(–4) layers of alar cells.
In Europe so far reported only from Baden-Württemberg in Germany (ENGELMARK 1999,
SAUER 2000), and recently discovered in Switzerland based on a collection from 1984 [Ct.
Bern, Simmental, Erlenbach, Stockenfluh, 1940 m a.s.l., I. Bisang 84246 (S; B78569)]. We
therefore believe that the species most likely occurs also in other localities in Central Europe.
– Grows in calcareous, semi-shady, dry habitats, on humus-covered rocks as well as on hu-
mus-rich stony ground (SAUER 2000).
Fig. 37: Dicranum brevifolium,
transverse section of leaf lamina
(NYHOLM 1987).

HEDENÄS & BISANG: Key to European Dicranum species 189
Dicranum drummondii Müll. Hal. (D. elatum Lindb.;
D. robustum Blytt ex B., S. & G.)
D. drummondii belongs to the large-sized Dicranum
species, comparative in size to D. majus or D.
polysetum. Its leaves have an ovate base and nar-
row gradually towards a long, falcate acumen (fig.
38). The upper leaf lamina is rugose, and has got
distinct dorsal, spine-like or highly conical projec-
tions, often sitting in transverse or oblique rows.
This species occurs in the northern parts of Europe.
– It is a characteristic species in dry pine forest, ei-
ther on dry sandy soils or on rocky ground.
Dicranum elongatum Schleich. ex Schwägr. [D.
sendtneri Limpr., p.p.; D. elongatum subsp. sendtneri (Limpr.) Podp., p.p.]
D. elongatum characteristically grows in very dense cushions (fig. 39a), usually so dense that
it is difficult to push your fingers into a cushion. The leaves are relatively narrow throughout,
when moist erect or almost so and straight or somewhat falcate, when dry slightly crisped, the
costa is strong, and both the upper and lower (fig. 39c) leaf lamina cells are strongly incrassate.
D. groenlandicum also forms very dense cushions, but can be differentiated from D. elongatum
by having long, porose cells throughout the leaves, whereas D. elongatum has short, eporose
cells in the upper portion of the leaf. While the leaf point is sharp to blunt in D. elongatum (fig.
39b), it is blunt to rounded in D. groenlandicum.
LIMPRICHT (1886) described D. sendtneri from the Sudetes. According to the original de-
scription, D. sendtneri should differ from D. elongatum in having longer and more curved
leaves, denticulate leaf margins and costae (supposedly entire in D. elongatum), a narrower
costa (
1
/
7
–
1
/
6
vs.
1
/
4
–
1
/
3
of maximal basal leaf width), and elongate upper leaf lamina cells
(quadrate in D. elongatum). We studied S material of D. elongatum and such that should
belong to D. sendtneri according to the labels, but were not able to recognise two different
taxa. Moreover, the isotype material of D. sendtneri Limpr. that is present in S consists
partly of D. elongatum and partly of
D. fuscescens (annotated by T.-B.
Engelmark).
D. elongatum is widespread in north-
ern and mountainous parts of Europe.
– It grows on more or less acid soil,
on heaths, in bogs or in bog portions
of poor mires, and on rocks.
Dicranum flagellare Hedw. [Ortho-
dicranum flagellare (Hedw.) Loeske]
D. flagellare is usually easily recog-
nised by its small size in combination
with the almost universal occurrence
of numerous flagellae that arise in the
upper leaf axils.
Fig. 38: Dicranum drummondii, dry shoots.
a b c
Fig. 39: Dicranum elongatum. a – dry shoots, b – leaf apex, c –
basal lamina cells.

190 Herzogia 17 (2004)
Widespread in Europe, rare or absent in the far north. – Mostly found on decomposing wood,
humus-rich soil, or peat, in more or less humid habitats.
Dicranum flexicaule Brid. [D. congestum var. flexicaule (Brid.) Bruch & Schimp.; D.
congestum auct.; D. fuscescens subsp. congestum auct. (GEISSLER et al. 1998)]
D. flexicaule is very similar to D. fuscescens, but has more irregular upper leaf lamina cells than the
latter, and less regularly serrate leaf margins and back of costa, the latter being serrate in at most the
upper third of the leaf. The differences between these two species are discussed under D. fuscescens.
Widespread in Europe, but apparently rarer in the south than in the north. – Grows on soil,
rocks, stumps, and logs, from the lowlands to high altitudes in the mountains.
Dicranum fragilifolium Lindb. [Orthodicranum fragilifolium (Lindb.) Podp.]
D. fragilifolium is one of the three European Dicranum species with straight and erect, stiff,
and fragile leaves. It differs from D. tauricum in having stereid bands in the basal part of the
costa, in having porose basal leaf lamina cells, and at least partly bistratose alar cells. It can be
separated from D. viride by its undifferentiated costal epidermal cells, its longer and porose
basal lamina cells, and its unistratose leaf lamina. It differs from both the other species in
having an asymmetrical and more or less curved spore capsule.
This species is widespread in the northern parts of Europe. NYHOLM (1987) reported D.
fragilifolium from the mountains of C Europe, but this could not be confirmed by additional
literature reports. – It grows on logs, stumps, and humus-rich soil, mostly in humid habitats.
Dicranum fulvum Hook. [Paraleucobryum fulvum (Hook.) Loeske]
D. fulvum forms dark green or sometimes brownish tufts. The leaves are falcate when moist and
curled when dry, and give a somewhat rigid impression. The leaf acumen may be slightly fragile.
This species is sometimes confused with D. scottianum or D. viride, but the leaf margin is
denticulate far down in D. fulvum (fig. 40a), almost entire or with a few denticles close to the
apex in the other two. The basal lamina cells are mainly rectangular in D. fulvum and D. viride
(fig. 40b), but predominantly or in large portions linear in D. scottianum. Finally, the leaves are
straighter and stiffer in D. scottianum and D. viride than in D. fulvum. D. fulvum could also be
confused with D. fuscescens, but can be separated from the latter by its broader and more longly
excurrent costa, larger portions of leaf lamina bistratose, its stiffer and sometimes fragile leaves,
and its more strongly incrassate lamina cells.
This species is widespread in Europe, except in the far north. – Found
on usually shaded (except in strongly humid environments) siliceous
rocks, rarely also on stem bases, mostly in deciduous, but sometimes
also in coniferous forest.
Dicranum fuscescens Turner [D. congestum Brid.; D. fuscescens
subsp. congestum (Brid.) Kindb.; D. sendtneri Limpr., p.p.; D.
elongatum subsp. sendtneri (Limpr.) Podp., p.p.]
D. fuscescens is a medium-sized Dicranum species (see fig. 43, under D.
majus) with narrow upper portions of the leaves, which are more or less
secund when moist and crisped when dry. The back of the costa is densely
serrate or mammillose in the upper
2
/
5
–
4
/
5
and the leaf margin is denticu-
late in the upper
1
/
3
–
1
/
2
.
a b
Fig. 40: Dicranum fulvum.
a – upper leaf acumen,
b – basal leaf lamina cells.

HEDENÄS & BISANG: Key to European Dicranum species 191
D. fuscescens is often confused with D. flexicaule, which has more irregularly shaped
upper lamina cells, and which leaf margins and costa back are less regularly serrate, the
latter at most in upper c.
1
/
3
of the leaf. However, even if extreme phenotypes are readily
distinguished from each other, other phenotypes appear almost intermediate and it is not
clear to us whether these two taxa are really good species. Some characters that suppos-
edly separate them, such as, whether the spore capsule is furrowed when dry and empty
(= D. fuscescens) or not, and the differences in leaf curvature and plant size, seem unreli-
able. According to observations of L.H., the states of the capsule are not correlated with
the gametophytic characteristics that supposedly define these two taxa. D. fuscescens
could also be confused with D. fulvum; characters separating these two are discussed
under the latter. Regarding the identity of D. sendtneri Limpr., see the note under D.
elongatum.
Widespread in Europe, but seemingly rarer than D. flexicaule in the north. – Found on soil,
rocks, stumps, and logs, from the lowlands to the alpine zone.
Dicranum groenlandicum Brid. [D. elongatum subsp. groenlandicum (Brid.) Mönk.]
D. groenlandicum grows in very dense cushions (fig. 41a). The upper portions of the leaves
are tubular, the leaf point is blunt to rounded, the upper lamina cells are prosenchymatous, and
both the lower and upper lamina cells are strongly incrassate and porose (figs 41b–d). In these
features, D. groenlandicum is similar to D. laevidens, but the latter species grows in loose
tufts (fig. 42a). The differences between D. groenlandicum and D. elongatum, which also
grows in very dense cushions, are discussed under the latter species.
The distribution of this species is confined to northernmost Europe. – Grows on moist or
periodically wet soil in tundra vegetation, or in bogs.
Dicranum laevidens R. S. Williams
D. laevidens has tubular upper leaf portions, the leaf point is blunt to rounded (fig. 42b), the
upper lamina cells are prosenchymatous, and both the lower and upper lamina cells are strongly
incrassate and porose. In these features, D. groenlandicum is similar to D. laevidens, but the
present species grows in loose tufts (fig. 42a).
Only known from northernmost Europe. – Found in intermediate to rich fens, in relatively dry
places in the fens.
a b c d a b
Fig. 41: Dicranum groenlandicum. a – habit (moist), b – ba- Fig. 42: Dicranum laevidens. a – habit (moist),
sal leaf lamina cells, c – upper leaf lamina cells, d – upper b – upper part of leaf acumen.
part of leaf acumen.

192 Herzogia 17 (2004)
Dicranum leioneuron Kindb. (D. bonjeanii
var. anomalum C. E. O. Jensen; D. bonjeanii
var. integrifolium H. Lindb.)
D. leioneuron belongs to the species having
prosenchymatous upper leaf lamina cells.
It is almost invariably found with flagellae
arising from the apex of the shoots, and
has entire or almost entire leaf margins. It
is sometimes confused with D. bonjeanii,
and the characters separating these species
are given under the latter.
D. leioneuron has been reported both from
N and C Europe, although its exact geographical distribution is unclear. – Found both in bogs, among
Sphagnum, in poor fens and in areas with high rainfall occasionally in more xeric habitats in rock
crevices or on boulders (CORLEY 1991).
Dicranum majus Turner
When well developed, D. majus is one of the largest members of the genus (fig. 43), with
regularly falcate-secund leaves. Its leaves are gradually narrowed to a long, fine point, and the
upper lamina cells are prosenchymatous and often partly bistratose shortly inside the leaf
margin. In the alpine region of the mountains and in the Arctic the plants are frequently smaller
and the leaves less distinctly falcate. Such plants may be confused with D. scoparium that,
however, has got distinct ridges on the upper back of the costa rather than the furrows and
serrations found in D. majus. In transverse section most of the upper leaf lamina cells are
rectangular in D. majus, quadrate in D. scoparium, and when sporophytes are present there
are 1–5 per perichaetium in D. majus, but only one in D. scoparium. D. majus, as presently
understood, is a variable taxon that still needs to be investigated in more detail. Characters
that are variable include the number of layers of guide cells in the costa, leaf curvature and
length, whether the upper leaf lamina is partly bistratose or not, and whether or not spinosely
projecting cell ends occur in the upper lamina.
Widespread in Europe. – Occurs both on humid to wet soil and on rocks, in forest and in more
open habitats, the latter especially in mountains and in the far north.
Dicranum montanum Hedw. [Orthodicranum montanum (Hedw.) Loeske]
D. montanum belongs to the small species of the genus (fig. 44). Its leaves are strongly crisped
and curled when dry. In habit it looks more like Dicranoweisia cirrata (Hedw.) Lindb. than it
resembles other Dicranum species. From Dicranoweisia cirrata it can be differentiated by its
denticulate rather than entire leaf margin, and by its differentiated alar cells. – We have seen a
specimen of D. cf. montanum which deviates from typical D. montanum in having a partly
bistratose upper leaf lamina and fragile leaves.
Widespread in Europe. – On rotten wood, roots, tree bases, tree stems
and rocks, more rarely on soil.
Dicranum muehlenbeckii Bruch & Schimp.
D. muehlenbeckii has tubular leaves that are slightly contorted when
moist and strongly curled when dry. The upper leaf lamina cells are
relatively thin-walled, and the margin is serrate close to the leaf apex.
Fig. 43: Dicranum fuscescens (left) and D. majus, habit (al-
most dry).
Fig. 44: Dicranum
montanum, habit (dry).

HEDENÄS & BISANG: Key to European Dicranum species 193
This species was often confused with D. brevifolium earlier, and the
differences between these species are mentioned in the note under the
latter.
Occurs in C Europe, and is in addition known from one locality in
central Sweden. – Grows on soil, rocks, occasionally on rotten wood,
in calcareous or mineral-rich and dry environments.
Dicranum polysetum Sw. (D. rugosum Brid.; D. undulatum F. Weber
& D. Mohr)
D. polysetum belongs to the large species in the genus. It is usually
easily recognised already in the field because of its typical leaf orien-
tation (fig. 45), with the uppermost leaves erect and those further down
spreading. The leaves are in addition transversely undulate and the leaf margin is sharply
dentate above. Although SMITH (1978) states that the margin is recurved below, it is actually
broadly incurved.
D. polysetum occurs all over Europe. – It grows on soil, rocks, and rotten wood, both in dry
habitats, such as pine heath forest, and in wet places such as swampy forest or tree-covered
bogs.
Dicranum scoparium Hedw. (D. tectorum Warnst. & H. Klinggr.)
D. scoparium belongs to the medium-sized species of the genus. In some respects it is one of
the most variable Dicranum species (fig. 46); both phenotypes with straight and falcate leaves
are frequent, and both dwarf and large male plants occur. The species has prosenchymatous
upper leaf lamina cells, and has in addition (usually four) distinct lamellae on the back of the
upper costa. Differences between D. scoparium, D. crassifolium, D. transsylvanicum, and poorly
developed plants of D. majus are discussed under the latter species. Plants from extreme dry or
Arctic environments may lack dorsal costal lamellae completely, and could therefore be con-
fused with D. bonjeanii. However, in D. scoparium the lower costa is strong, dorsally convex,
and has several stereid bands, whereas that of D. bonjeanii is weak, almost flat, and has fewer
stereid bands (NYHOLM 1987, FRISVOLL & ELVEBAKK 1996).
Widespread in Europe. – Found on various substrates, on soil, rock, rotten wood, tree stems
etc., in both dry and humid environments.
Fig. 45: Dicranum
polysetum, moist shoot.
Fig. 46: Dicranum scoparium, examples of habit variation (moist). a – plants with straight leaves from pine heath
forest; b – plants with falcate leaves from swampy forest; c – large plants with falcate leaves from high-altitude
laurel forest on Madeira.
a b
c

194 Herzogia 17 (2004)
Dicranum scottianum Turner ex Scott, Robert [D. scottianum subsp. anglicum
Reimers; Orthodicranum scottianum (Turner ex Scott, Robert) G. Roth]
D. scottianum is dark green, and has rigid, hardly falcate leaves that are slightly
curled when dry. The lamina has bistratose margins and the basal lamina cells
are predominantly elongate (fig. 47). For differences between this species, D.
canariense and D. fulvum, see the notes under the latter two.
As shown by NELSON (1997), the correct author citation for D. scottianum is
‘Turner ex Scott, Robert’.
D. scottianum occurs in oceanic areas of Europe, and in addition in the Ca-
nary Islands and the Azores. Previously reported from Switzerland (URMI et
al. 1997), but the relevant specimen (Ct. Bern, Interlaken, 1855, Bamberger,
S) belongs to D. fulvum. – Grows on siliceous rocks, mostly in the vicinity of
the seacoast.
Dicranum spadiceum J. E. Zetterst. [D. muehlenbeckii var. spadiceum (J. E. Zetterst.) Podp.;
D. neglectum Jur. ex De Not.]
D. spadiceum belongs to the medium-sized species of the genus. Its leaves are distinctly tubular
and the leaf margin is entire or in the upper part somewhat denticulate. The uppermost lamina
cells are irregular, with rounded corners (fig. 48c); further down the cells gradually become
more elongate and porose (fig. 48b). Sometimes the non-porose lamina cells are restricted to the
very uppermost lamina portion. When dry, the leaves are erect-spreading and slightly crisped
(fig. 48a).
This species has sometimes been confused with D. muehlenbeckii, which leaves are more strongly
curled when dry, and which margins are serrate near the apex. Leaf lamina cell size (cf. NYHOLM
1987) seems to be an unreliable distinguishing character between these two species.
Widespread in mountainous areas in Europe, but probably overlooked in many areas, espe-
cially in Central Europe. – Most commonly found on moist soil or rocks, sometimes in shal-
low mountain mires or on rotten logs, usually at relatively high altitudes.
Dicranum spurium Hedw.
Dry plants of D. spurium are easily
recognised by their shoot apices,
which remind about lettuce heads.
Otherwise, the leaves have a rela-
tively broad basal portion, narrow
suddenly to a short acumen, and the
upper lamina cells are very irregu-
lar (fig. 49). On the back of the up-
per lamina scattered or numerous
cells have usually got conical mam-
millae.
This species is widespread in Europe,
rare or absent in the far north. – Forms
dense colonies on soil or rock, in
more or less dry and open forests.
a b c
Fig. 48: Dicranum spadiceum. a – dry shoot; b – median leaf
lamina cells; c – upper leaf lamina cells.
Fig. 47: Dicra-
num scottianum,
basal leaf lamina
cells.

HEDENÄS & BISANG: Key to European Dicranum species 195
Dicranum tauricum Sapjegin [D. strictum Schleich. ex D. Mohr;
Orthodicranum strictum Broth.; O. tauricum (Sapjegin) Smirnova]
D. tauricum has straight and erect, stiff and fragile, upper leaves. It
differs from the other two such species, D. fragilifolium and D. viride,
in lacking stereid bands in the costa. From D. viride it also differs in its
unistratose leaf lamina. Additional differences between this species and
D. fragilifolium are discussed under the latter.
This species is widespread in W and C Europe, and appears to have
increased in many areas, possibly due to acidification. – Occurs on tree
bases and rotten wood, rarely on soil.
Dicranum transsylvanicum Lüth
D. transsylvanicum was recently described from Romania, and the follow-
ing information is taken from the original description (LÜTH 2002). Like
D. crassifolium, this species has a partly bistratose upper leaf lamina and
lamellae on the upper back of the costa. However, contrary to both D.
crassifolium and D. scoparium, which remind about the present species in
having well-developed lamellae on the back of the costa, D. transsylvanicum
has a strongly spinose dorsal lamina and irregularly spinose-dentate upper
leaf margins (fig. 50). D. scoparium has in addition a unistratose leaf lamina.
Only known from the Apuseni Mountains in western Romania. – The
single known specimen was collected on spruce needle litter between
limestone boulders, at around 1150 m a.s.l.
Dicranum undulatum Schrad. ex Brid. (D. affine Funck; D. bergeri
Bland.; D. schraderi Wahlenb.)
D. undulatum grows in rather dense tufts. In addition, it is recognised by its straight or at most
weakly curved leaves (fig. 51a), that are more or less distinctly transversely undulate, lanceolate,
and narrow very gradually upwards, except that they are suddenly narrowed to the obtuse apex in
Fig. 50: Dicranum trans-
sylvanicum, portion of
upper leaf lamina and
costa seen from dorsal
side (after LÜTH 2002).
Fig. 51: Dicranum undulatum. a – habit (moist), b – portion of dry shoot, c – cells of upper leaf lamina.
a b c
Fig. 49: Dicranum
spurium, cells of upper
leaf lamina.

196 Herzogia 17 (2004)
the uppermost part (fig. 51b). Although frequently called D. bergeri by modern European authors,
the correct name and citation for this species appears to be ‘D. undulatum Schrad. ex Brid., J. f.
Bot. 1800, 1(2): 294. 1801’. The latter name is thus a few years older than D. bergeri.
D. undulatum is widespread in Europe. – Occurs on moist or wet ground, usually on bogs or
hummocks in mineral-poor fens, sometimes also in meadows. In the northern boreal zone the
species is in addition frequent in dry pine heath forest.
Dicranum viride (Sull. & Lesq.) Lindb. [Paraleucobryum viride (Sull. & Lesq.) Podp.]
D. viride belongs to the Dicranum species with straight and erect, stiff, and fragile leaves,
among which it is characterised by its partly bistratose leaf lamina. According to NYHOLM
(1987) and observations by ERZBERGER (1999) on Hungarian material, D. viride differs from
D. tauricum and D. fragilifolium by cell wall projections which are visible in leaf cross sec-
tions (compare D. brevifolium; fig. 37). Additional differences between this species, D.
fragilifolium, and D. tauricum are discussed under the latter two.
This species occurs in Central Europe, northwards to southern Norway and the Baltic Sea
area. – Mostly found as an epiphyte on stems of deciduous trees, more rarely on coniferous
trees, rocks, or soil.
Acknowledgements
We thank the participants of the Dicranum workshop in Zürich in April 2003, M. Sauer, and M. Siegel for sugges-
tions that improved the key, and the “Schweizerische Akademie der Naturwissenschaften”, SANW, for supporting
the travel costs in connection with the workshop.
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Bundesamt für Umwelt, Wald und Landschaft (BUWAL), Schriftenreihe Umwelt 265: 1–47.
Manuscript accepted: 13 February 2004.
Addresses of the authors
Lars Hedenäs & Irene Bisang, Swedish Museum of Natural History, Department of Crypto-
gamic Botany, Box 50007, SE-104 05 Stockholm, Sweden.
E-mail: lars.hedenas@nrm.se, irene.bisang@nrm.se
Species index
Dicranoweisia
cirrata 192
Dicranum
acutifolium 185, 186, 187, 188
affine 195
amannii 187
angustum 180, 186, 187
bergeri 195, 196
var. acutifolium 187
bonjeanii 180, 183, 187, 192, 193
var. anomalum 192
var. integrifolium 192
brevifolium 181, 184, 186, 187, 188, 193, 196
canariense 180, 184, 185, 188, 194
congestum 190
auct. 190
var. flexicaule 190
crassifolium 183, 188, 193, 195
dispersum 181, 184, 186, 187, 188
drummondii 180, 183, 189
elatum 189
elongatum 180, 186, 189, 191
subsp. groenlandicum 191
subsp. sendtneri 189, 190
flagellare 180, 181, 184, 189
flexicaule 180, 185, 190, 191
fragilifolium 180, 182, 190, 195, 196
fulvum 180, 184, 190, 191, 194
fuscescens 180, 181, 185, 189, 190, 191, 192
subsp. congestum 190
subsp. congestum of Geissler et al. 190
groenlandicum 180, 182, 189, 191
laevidens 180, 182, 191
latifolium 187
leioneuron 180, 182, 187, 192
majus 180, 181, 183, 189, 190, 192, 193
montanum 180, 181, 184, 192
muehlenbeckii 186, 188, 192, 194
auct. 188
var. acutifolium 187
var. brevifolium 188
var. spadiceum 194
neglectum 194
palustre 187
polysetum 180, 181, 183, 189, 193
robustum 189
rugosum 193
schraderi 195
scoparium 180, 181, 183, 187, 188, 192, 193, 195
scottianum 180, 181, 184, 185, 188, 190, 194
subsp. anglicum 194
sendtneri 189, 190, 191
spadiceum 182, 186, 194
spurium 180, 183, 194, 195
strictum 195
tauricum 180, 181,182, 190, 195, 196
tectorum 193
transsylvanicum 183, 188, 193, 195
undulatum 181, 183, 195, 196
undulatum Turner 187
undulatum F. Weber & D. Mohr 193
viride 180, 181, 182, 190, 195, 196
Kiaeria
glacialis 180
Orthodicranum
flagellare 189
fragilifolium 190
montanum 192
scottianum 194
strictum 195
tauricum 195
Paraleucobryum
fulvum 190
viride 196