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Arctoa (1996) 6: 21-112
BRYOPHYTE FLORA OF ALTAI MOUNTAINS. VII. HYPNACEAE AND
RELATED PLEUROCARPS WITH BI- OR ECOSTATE LEAVES
ÌÎÕÎÎÁÐÀÇÍÛÅ ÀËÒÀß. VII. HYPNACEAE È ÁËÈÇÈÅ ÑÅÌÅÉÑÒÂÀ
ÁÎÊÎÏËÎÄÍÛÕ ÌÕÎÂ Ñ ËÈÑÒÜßÌÈ ÁÅÇ ÈËÈ Ñ ÄÂÎÉÍÎÉ ÆÈËÊÎÉ
MICHAEL S. IGNATOV1, HISATSUGU ANDO2 & ELENA A. IGNATOVA3
ÌÈÕÀÈË Ñ. ÈÃÍÀÒÎÂ1, ÕÈÑÀÑÒÓÃÓ ÀÍÄÎ2 È ÅËÅÍÀ À. ÈÃÍÀÒÎÂÀ3
Abstract
The family Hypnaceae in Altai includes the genera Callicladium (1 species), Eurohypnum
(1), Gollania (1), Heterophyllium (1), Homomallium (2), Hypnum (11), Podperaea (1),
Ptilium (1), Pylaisiella (2), Taxiphyllum (1); family Plagiotheciaceae – Herzogiella (1),
Isopterygiopsis (3), Myurella (3), Orthothecium (3), Plagiothecium (4), Platydictya (1);
family Entodontaceae – Entodon (3), Platygyrium (1); family Hylocomiaceae –
Hylocomiastrum (2), Hylocomium (1), Pleurozium (1), Rhytidiadelphus (2); family
Sematophyllaceae – Struckia (1). Descriptions and illustrations are given for all species.
Homomallium connexum and Hypnum fauriei are reported in South Siberia for the first
time. Some evidence for placement of Platygyrium in the Entodontaceae is provided.
Abstract
Ñåìåéñòâî Hypnaceae íà Àëòàå âêëþ÷àåò ðîäû Callicladium (1 âèä), Eurohypnum
(1), Gollania (1), Heterophyllium (1), Homomallium (2), Hypnum (11), Podperaea (1),
Ptilium (1), Pylaisiella (2), Taxi phyllum (1); ñåìåéñòâî Plagiotheciaceae – Herzogiella
(1), Isopterygiopsis (3), Myurella (3), Orthothecium (3), Plagiothecium (4), Platydictya
(1); ñåìåéñòâî Entodontaceae – Entodon (3), Platygyrium (1); ñåìåéñòâî Hylocomiaceae
– Hynocomiastrum (2), Hylocomium (1), Pleurozium (1), Rhytidiadelphus (2); ñåìåéñòâî
Sematophyllaceae – Struckia (1). Äëÿ âñåõ âèäîâ äàíû îïèñàíèÿ è èëëþñòðàöèè.
Homomallium connexum è Hypnum fauriei óêàçûâàþòñÿ äëÿ Þæíîé Ñèáèðè âïåðâûå.
Ïðèâåäåíû àðãóìåíòû â ïîëüçó ïåðåâåäåíèÿ Platygyrium â ñåìåéñòâî Entodontaceae.
1 – Main Botanical Garden of Russian Academy of Sciences, Botanicheskaya 4, Moscow 127276 Russia – Ðîññèÿ 127276
Ìîñêâà, Áîòàíè÷åñêàÿ 4, Ãëàâíûé áîòàíè÷åñêèé ñàä ÐÀÍ
2 – Department of Biological Science, Faculty of Science, Hiroshima University, Kagami-yama, Higashi-hiroshima,
Hiroshim-ken 739, Japan
3 – Department of Geobotany, Biological Faculty, Moscow State University, Moscow 119899 Russia – Ðîññèÿ 119899,
Ìîñêâà, Ìîñêîâñêèé óíèâåðñèòåò, Áèîëîãè÷åñêèé ôàêóëüòåò, êàô. ãåîáîòàíèêè
This paper continues the series on a descrip-
tive bryophyte flora of Altai Mts. started in the
third volume of Arctoa. A brief overview of the
environments of Altai and the history of ex-
ploration of its moss flora have been published
by Ignatov (1994).
In preparing this paper, the authors worked
together on identification and description of
species. However discussion on generic/famil-
ial classification are solely the responsibility of
Ignatov, whose ideas sometimes disagree with
those of Ando (Ando, 1972, 1973; Nishimura &
al., 1984).
The Hypnaceae are one of the families of
mosses poorly delimited from the related fam-
ilies, Sematophyllaceae, Plagiotheciaceae,
Entodontaceae, and Hylocomiaceae, and its cir-
cumscription is treated variously by different
authors. Recent publications contribute much
to this topic, resulting in suggestions of chang-
ing of familial position for many genera. How-
ever some of these new ideas are still in dispute,
so it seems better to describe species of these
families in one paper.
The following genera are included in this
paper:
Hypnaceae: Callicladium, Eurohypnum, Gol-
lania, Heterophyllium, Homomallium, Hypnum,
Podperaea, Ptilium, Pylaisiella, Taxiphyllum;
Plagiotheciaceae: Herzogiella, Isopterygiop-
22 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
sis, Myurella, Orthothecium, Plagiothecium, Pla-
tydictya;
Entodontaceae: Entodon, Platygyrium;
Hylocomiaceae: Hylocomiastrum, Hylocomi-
um, Pleurozium, Rhytidiadelphus;
Sematophyllaceae: Struckia.
Definition of the Amblystegiaceae also pos-
es some problems around Calliergonella, Campy-
lium, Campyliadelphus, Campylophyllum, and
Pseudohygrohypnum. These genera will be dis-
cussed in the paper on Amblystegiaceae. How-
ever some of them are included in the key of
the genera (marked by an asterisk rather than
page).
The terminology used in the paper is mostly
standard. However some explanations are nec-
essary.
(1) Pseudoparaphyllia. Though this organ
was specially discussed by some authors, the
difference between pseudoparaphyllia in a strict
sense and juvenille branch leaves remains dis-
putable. Enroth (1994) suggested considering
pseudoparaphyllia as foliose organs, usually lan-
ceolate, situated obviously apart from branch
initial and not confused with broadly ovate-
triangular juvenile branch leaves. According to
this conception pseudoparaphyllia occur in rel-
atively few species, while most of the species
have no pseudoparaphyllia. Hedenäs (1995)
used the term pseudoparaphyllia for both lan-
ceolate structures around and broad structures
at the base of juvenile branches. He considered
pseudoparaphyllia absence a rare phenomenon
when after branch removal nothing leaf-like
remains on stem (as in Plagiotheciaceae). We
use here the latter teminology. One problematic
case in pseudoparaphyllia interpretation is dis-
cussed under Orthothecium.
(2) Rhizoid position. In most pleurocarps,
rhizoids are clustered on the stem just below
the leaf insertion. In the zone of close contact
with the substrate, rhizoids are usually more
numerous and in part may arise from the abax-
ial side of leaves. However, in places where
few rhizoids are present their position is clear-
ly cauline, and typically they form a trans-
verse row or a transversely ovoid group. In
some genera under question rhizoids are axil-
lary. This means their position is in leaf axils,
usually 1-2 cells above the leaf. Axillary rhiz-
oids are usually fewer, purplish, finely granu-
lose and form a longitudinal row or other-
wise longitudinally ovoid group (if rhizoids
are two, they are nearly always one above an-
other). Some deviation from this type is seen
in Herzogiella where rhizoids are shifted more
distally from the leaf below them and grouped
mostly near the corner of the leaf situated
above; however this type is considered pri-
marily axillary due to rhizoid texture, often
longitudinally seriate positioning and also
sometimes a more clearly axillary pattern. The
similar position of axillary rhizoids is ob-
served at branch tips of Hylocomiaceae,
Entodon, Ptilium. In Plagiothecium rhizoids
are confined to the abaxial side of leaf, most-
ly along the costa, and this pattern was seen
also in branches of some specimens of Entodon.
In proximal branches of Plagiothecium rhiz-
oids are numerous and sometimes partly
cauline, but the places with dense rhizoids seem
too abnormal to take them into consideration.
Rhizoid position of Struckia is discussed
under that genus. Rhizoids on branches often
have a different position than those on stems
and also are variable even within a single
species (e. g., Entodon concinnus).
(3) Leaf length of falcate leaves indicate
the approximate length if they were straight;
although there is no precise procedure of mea-
surements in this case, we think that without
such an approximation, the description of spe-
cies looks incomplete.
(4) Laminal cell width is measured as the
width of lumen + cell wall, with an indication
in brackets of widths of the two, respectively.
(5) Capsule length is measured in dry ino-
perculate herbarium material.
(6) Annulus is described according Buck
(1980), only for a structure with separating cells.
The situation when a deciduous annulus is not
seen at the urn mouth is described as annulus
none, though in fact thick-walled hygroscopic
cells remain attached to the operculum or to
the urn.
(7) The terms lamella and trabecula were
both applied to the projecting walls on either
surface of the exostome tooth. We will name them
dorsal and ventral trabeculae. Papillae on the
dorsal trabeculae are usually very similar to those
on dorsal vertical projections, so the descrip-
tion of papillae on the dorsal trabeculae implies
23
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
also papillae on the dorsal vertical projections.
Ventral trabeculae in the proximal part of tooth
are smooth (non-papillose) always, above the
transition zone are papillose or smooth, and in
the uppermost part of tooth are smooth again
in many species which have papillose ventral
trabecule in the middle. In description smooth
or papillose ventral trabeculae mean the state
of this character at about 3/4–4/5 of tooth
length.
(8) On SEM micrographs the margin of the
exostome tooth in its distal portion is often
differentiated, becoming more thick. We call this
margin incrassate (figs. on pp. 104, 110, etc.) or
non-incrassate (figs. on pp. 47, 49, etc.).
SEM observations were made with Hitachi
S-405A SEM at 15 kV. For these observations
were taken dry capsules from herbarium, most-
ly from Altaian collection. Peristomes were coat-
ed by gold without additional preparation.
KEY TO THE ALTAIAN GENERA OF
HYPNACEAE AND RELATED PLEUROCARPS
WITH BI- OR ECOSTATE LEAVES:
1. Paraphyllia numerous, branched, densely cov-
ering the stem; robust mosses . . . . . . . . . . . 2
1. Paraphyllia absent or rarely simple, lan-
ceolate, not conspicuous . . . . . . . . . . . . . . . . 3
2. Costa double and short; stem regularly
bipinnate-branched; plants each year form-
ing flat fronds resulting in stepwise ap-
pearance . . . . . . . . . Hylocomium (p. 104)
2. Costa single or double, to mid-leaf or long-
er; plants less regularly 1-2(3)-pinnate-
branched, without stepwise appearance
. . . . . . . . . . . . . . . Hylocomiastrum (p. 99)
3. Leaves broadly obtuse; robust mosses with
stems ascending to erect . . . . . . . . . . . . . . 4
3. Leaves acute to acuminate, usually ±pros-
trate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4. Stem deep-red . . . Pleurozium (p. 107)
4. Stem green to yellow-brown . . 5
5. Alar cells large, inflated, forming a well-de-
limited group . . Calliergonella cuspidata*
5. Alar cells ±small . . Entodon concinnus (p. 25)
6. Leaves strongly reflexed (or sometimes
spreading at right angle from the base) . . 7
6. Leaves not reflexed . . . . . . . . . . . . . . . . 10
7. Plants robust . . . . . . . . . . . . . . . . . . . . . . . . . 8
7. Plants rather small . . . . . . . . . . . . . . . . . . 9
8. Leaves serrate, auriculate; largest cells at
base in the middle between the costa and
margin; costa double, prominent, to 1/3-
2/3 of the leaf length
. . . . . . . . . . Rhytidiadelphus (p. 105)
8. Leaves ±entire, not auriculate; largest cells
at base in leaf corners; costa double, usu-
ally short (rarely longer, up to 1/3 the
leaf length)
. . . Campylium p. p. (Campyliadelphus)*
9. Teeth in lower leaf margins formed by two
cells: upper end of the lower cell and lower
end of the upper cell; laminal cells distinct-
ly prorate . . . . . . . . . . . . . Podperaea (p. 60)
9. Teeth along the whole leaf margin formed
by one cell; laminal cells smooth
. . . . . . . Campylium (Campylium s. str. &
Campylophyllum)*
10.Leaves decurrent; rhizoids abaxial near
leaf bases; plants medium-sized to small,
rarely branched, ±complanately foliate
. . . . . . . . . . . . . . . . . Plagiothecium (p. 46)
10.Leaves non-decurrent; rhizoid position
otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11.Rhizoids on the stem clearly axillary, 1-2
cells above the leaf axil (rarely higher above
the axil in Herzogiella), usually purple and
finely granulose . . . . . . . . . . . . . . . . . . . . . . . 12
11.Rhizoids on the stem non-axillary, appear-
ing mostly below the leaf base (but some-
times ±axillary at branch tips), usually
brown and smooth . . . . . . . . . . . . . . . . . . . 16
12.Plants small, imbricate, very fragile; leaves
strongly concave, short, usually with length
to width ratio less than 2:1; laminal cells
prorate or papillose on the abaxial sur-
face . . . . . . . . . . . . . . . . . . Myurella (p. 40)
12.Plants otherwise . . . . . . . . . . . . . . . . . . . 13
13.Plants small to robust, ±orthotropic, often
tinged with bronze or red; leaves straight,
acuminate; on moist rock outcrops or on soil
in tundra . . . . . . . . . . Orthothecium (p. 42)
13.Plants prostrate, green to yellow green . . 14
14.Plants medium-sized; leaves falcate, rath-
er loosely arranged, sharply serrate above,
serrulate downward to the base; capsules
slightly curved; rhizoids mostly situated
between leaves; epixylic autoicous plant
usually with sporophytes
. . . . . . . . . . . . . . . . Herzogiella (p. 53)
24 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
14.Plants otherwise; leaves less serrate,
smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15.Plants small to medium-sized, usually pure
green; epidermal cells larger or as large as
cells of the subtending layer; laminal cells
linear to short, 1:4-7 (I. alpicola); stem with
central strand . . . Isopterygiopsis (p. 35)
15.Very small thin green plants with loosely
arranged leaves; laminal cells short, 1:3-4;
stem without central strand
. . . . . . . . . . . . . . . . . . . . . . Platydictya (p. 39)
16.Leaves near shoot tips caducous, result-
ing in a subterminal naked zone on many
shoots; leaves laxly areolate
. . . . . . . . . . . . . . . . . . . . . . . . Struckia (p. 97)
16.Plants without leafless zones near shoot
tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . 17
17.Capsules perfectly straight; leaves symmet-
ric or nearly so . . . . . . . . . . . . . . . . . . . . . .. . 18
17.Capsules curved or/and somewhat inclined,
or plants without sporophytes; leaves fal-
cate-secund to ±symmetric . . . . . . . . . . . 20
18.Leaves ±shortly acute; plants distinctly
complanate . . . . . . . . . . . . Entodon (p. 25)
18.Leaves acuminate; plants ±julaceous . . 19
19.Plants dark, with brood branches clustered
in leaf axils, especially near branch ends; leaf
margins recurved . . . . . Platygyrium (p. 33)
19.Plants ±shiny, without brood branches; leaf
margins erect . . . . . . . . . Pylaisiella (p. 56)
20.Plants rather tall, erect, densely and reg-
ularly plumose-branched; leaves strong-
ly falcate to circinate, plicate
. . . . . . . . . . . . . . . . . . . . . . . . . . Ptilium (p. 91)
20.Plants more or less prostrate; leaves not
or scarcely plicate . . . . . . . . .. . . . . . . . . . 21
21.Plants complanate; leaves straight, acute to
shortly acuminate, serrulate; alar cells indis-
tinctly differentiated . . Taxiphyllum (p. 94)
21.Leaves longly, rarely shortly acuminate;
straight to falcate; alar cells ±differentiat-
ed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22.Leaves strongly curved (tip is turned
from 90° to 180°) . . Hypnum p.p. (p. 68)
22.Leaves symmetric or curved less than on
90° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23.Epidermal cells of stem enlarged, thin-
walled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
23.Epidermal cells of stem small, thick-walled
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24.Alar cells thin-walled, inflated
. . . . . . . . . . .. Calliergonella lindbergii *
24.Alar cells not inflated, ±opaque
. . . . . . . . . . . . . . . . . . .. . Hypnum p.p. (p. 68)
25.Autoicous, usually bearing sporophytes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
25.Dioicous, usually without sporophytes . . 28
26.Small plants, with ±straight leaves; usu-
ally on rocks . . . . . Homomallium (p. 63)
26.Large plants, usually on rotten wood (rare-
ly on trunks or rocks) . . . . . . . . . . . . . . 27
27.Plants golden-yellow; alar cells thick-walled
and brownish . . . . Heterophyllium (p. 90)
27.Plants green; alar cells thin-walled, hyaline-
transparent . . . . . . . . . . Callicladium (p. 86)
28.Leaves serrate in acumen, ±plicate; large
plants . . . . . . . . . . . . . . . . . Gollania (p. 92)
28. Leaves entire or with serrulate acumen . . 29
29.Leaves straight, concave, imbricate, distinct-
ly serrulate in upper part; alar regions ex-
tending 1/3 the leaf length
. . . . . . . . . . . . . . . . . . . . . . . Eurohypnum (p. 67)
29.Leaves ±falcate; alar cells not so numerous
. . . . . . . . . . . . . . . . . . . . . . . Hypnum p.p. (p. 68)
Entodontaceae Kindb.
The family was revised by Buck (1980), who
delimited it using mostly sporophytic charac-
ters, namely the reduction of endostome (basal
membrane low or lacking) and exostome (lack
of trabeculae on the inner surface of teeth). Most
species are autoicous (rarely dioicous), and have
terete or complanate foliage, straight capsules, a
tendency in ornamentation of the outer exos-
tome to form solid cristae, transverse below and
often oblique to longitudinal above.
Buck (1980) noted a peculiar ornamenta-
tion of the exostome in Platygyrium, though he
did not discuss it in detail. Ornamentation of
the outer exostome of Platygyrium is reticu-
late, but distinct from the reticulate pattern of
the Hylocomiaceae. In the latter it is a result of
fusing of transversal cristae, while in Platygy-
rium the main orientation of cristae is longitu-
dinal. Under the light microscope exostome teeth
of Platygyrium above the middle look papil-
lose, rather similar to those of most Hypnaceous
mosses. However the dots (“papillae”) comprise
25
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
an optical effect from centers of reticulum loops,
obvious from SEM micrographs (Figs. 000).
Buck (1980) retained Platygyrium in Hyp-
naceae, arguing this by the presence of normally
developed ventral trabeculae of the exostome
teeth. However high trabeculae on the inner sur-
face of teeth are known in some species of Entodon
(E. brevisetus (Hook. et Wilson) Jaeg., E. sca-
bridens Lindb., etc., cf. Mizushima, 1960)). In most
Entodon species they are absent in the distal
half of teeth but ±well-developed below. There-
fore a similarity in peristome structure between
Platygyrium repens and Entodon is more strik-
ing than with that of any genus of Hypnaceae,
and therefore Platygyrium is worth considering
as a member of the Entodontaceae.
Additional similarities of Entodon and
Platygyrium include (1) erect capsule; (2)
low conic operculum with long beak; (3) epi-
phytic and epixylic habitat preferences and
rather southern, hemiboreal to temperate distri-
bution (though some species of Hypnaceae also
have such ecology and distribution).
The gametophytic characters of Platygyrium
do not disagree with the Entodontaceae (mostly
because there are almost no valuable differences
in gametophytes of Hypnaceae and Entodonta-
ceae). The dioicous condition is unusual in
Entodontaceae, but some Entodon species are
dioicous, and the character of sexual condition is
not stable in most families of pleurocarps.
Entodon C. Müll.
The genus includes about 100 species with
predominantly tropical to temperate distribu-
tion. Only one species, Entodon concinnus, pen-
etrates the Arctic regions and high mountains
of the Northern Hemisphere. Several other spe-
cies, like Entodon cladorrhizans, E. schleicheri
and E. compressus, are principally temperate,
but also occur in the hemiboreal zone.
KEY TO THE ALTAIAN SPECIES OF ENTODON
1. Yellowish plants with ascending stems, dio-
icous; typically epigeic, rarely epilithic or ep-
iphytic; branches julaceous; leaves with broad-
ly rounded apex and non-transparent 2-3-stra-
tose alar parts . . . . . . . E. concinnus (p. 25)
1. Pure green plants with totally plagiotropic
stems, autoicous; typically epiphytic, occa-
sionally epilithic; stem and branches dis-
tinctly complanate; leaves acute to acumi-
nate at apex, with transparent unistratose alar
parts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Robust plants with shoots about 2 mm
broad at the tip; branches straight; seta
yellowish; capsule cylindric; exostome
teeth striolate below, papillose in the mid-
dle, weakly papillose to smooth above
. . . . . . . . . . . . . . . . . . . E. schleicheri (p. 28)
2. Medium-sized plants with shoots about
1 mm broad at the tip; branches distinctly
curved; seta reddish; capsule ovoid; exos-
tome teeth papillose below
. . . . . . . . . . . . . . . . . . . . E. compressus (p. 27)
Entodon concinnus (De Not.) Par., Index
Bryol., ed. 2, 2: 130. 1904. – Hypnum concin-
num De Not., Mem. Reale Accad. Sci. Torino 39:
220. 1836. – Entodon orthocarpus (Brid.)
Lindb., Musci Scand. 39. 1879. Fig. 1
Plants green to yellow-green, in loose mats. Stem
ascending to erect, to 10 cm long, with well-devel-
oped central strand, regularly pinnate-branching;
branches 10-13 mm long; branching often sympodi-
al, and in suboptimal conditions to 6 sympodial steps.
Pseudoparaphyllia broadly triangular. Axillary hairs
4-celled, 100 μm long, apical cell 38-50 × 10 μm; rhiz-
oids rare, on stem below leaf insertions, axillary on
branch tips or (on other plants) abaxial on leaves.
Stem leaves 1.9-2.6 × 1.0-1.3 mm, ovate, widest at
1/3 of leaf length, obtuse. Branch leaves 1.4-1.6 ×
0.7-0.8, widest at 1/2-2/3 of leaf length. Laminal
cells 75-115 × 5-6 (3-4.5/1.5-2.0) μm; alar cells nu-
merous, subquadrate, ca. 7 × 7 μm, thick-walled, form-
ing multi-stratose inflated areas.
Dioicous. Sporophytes found in Altai only in Bele
(mean annual temperature ca. +4°C, preci pitation 400
mm, rather open Larix + Betula forests), but repre-
sented only by old capsules with imperfectly pre-
served peristomes. Outer perichaetial leaves ±ob-
tuse, squarrose. Inner perichaetial leaves 3.5 mm long,
ecostate. Seta 1.5 cm. Capsule ca. 2.0 mm long. Exos-
tome teeth 0.35 mm long, densely papillose below,
±smooth above. Endostome segments of the same
length, ±smooth. Spores 18-21 μm.
The orthotropic julaceous habit is similar to
that of Pleurozium schreberi, but the latter spe-
cies differs at once by the red stem (easily seen
even through leaves).
Distribution: Entodon concinnus is widely dis-
tributed in the Holarctic, penetrating rather far to
the north in oceanic regions, including Iceland, Nor-
way and Sweden in Europe, Chukotka Peninsula,
Arctic Alaska, and Newfoundland. It is absent in
boreal lowlands, but common in mountains of both
26 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 1. Entodon concinnus (De Not.) Par. (1-13, 16-17 – from Yailyu 500 m Ignatov 0/996; 14-15, 18 – from Bele
500 m Ignatov 0/66): 1-2 – habit; 3-5 – stem leaves; 6 – upper laminal cells of stem leaf; 7-10 – branch leaves; 11
– stem cross-section; 12 – middle laminal cells of stem leaf; 13 – alar cells of stem leaf; 14-15 – perichaetial leaves;
16-17 – pseudoparaphyllia; 18 – part of peristome. Scale bars: 1 cm – for 1; 5 mm – for 2; 1 mm – for 3-10, 14-15;
0,2 mm – for 17; 100 μm – for 6, 11-13, 16, 18.
10
15
2
5
89
11
12
13
7
6
3
1
4
14
1 cm
5 mm
1 mm
100 μm
0.2 mm
17
16 18
Eurasia (Alpes, Carpatians, Caucasus, Pamir, Tian-Shan,
Altai, Sayan, mountains of Mongolia, Russian Far East,
China, Japan) and North America. Outside the Hol-
arctic it is known in New Guinea and Ecuador. In
the northern part of Altai E. concinnus has a rather
scattered distribution on rocks and sometimes on
fallen trunks, and rarely on living inclined trees in
the forest zone (not exceeding 1400 m). In drier areas
with forest-steppe vegetation E. concinnus becomes
more common, occurring in wider range of habitats:
over-grazed pastures (where it sometimes is the only
moss species), xerophytic shrubby steppes and form-
ing extensive carpets in Larix stands. In the most
xeric south-eastern part of Altai, on Kuraiskij Range
E. concinnus penetrates mountain tundra, reaching
2750 m, growing there on sheltered rock outcrops or
on soil bank along temporary stream beds.
Specimens examined: Aedigan Creek, 5 km upstream
600 m (34/44); Bele 500 m (0/66; 0/995; 0/997), 550 m
(0/71); Bolshoi Yaloman Creek, in middle course 1500 m
(Makarov 15.VII.1972); Chemal, 4 km upstream along
Katun 450 m (34/110); Chiri Creek, 0.5 km upstream 450
27
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 2. Entodon compressus C. Müll. (from Maima 280 m Ignatov 35/60): 1-2 – habit; 3 – capsule; 4-7 – stem
leaves; 8-9 – pseudoparaphyllia; 10 – upper laminal cells of stem leaf; 11-14 – branch leaves; 15 – stem cross-
section; 16-17 – perichaetial leaves; 18 – middle laminal cells of stem leaf; 19 – alar cells of stem leaf. Scale bars: 1
cm – for 1; 2 mm – for 2-3; 1 mm – for 4-7, 11-14, 16-17; 100 μm – for 8-10, 15, 18-19.
1
2
345
89
11 12 13 14
7
6
10
15
18 19
17
16
100 μm
1 mm
2 mm
1 cm
m (17/18); Elekmonar Creek, 5 km upstream 700 m (26/
101); Karagai 440 m (0/278), 450 m (Zolotukhin
11.VIII.1988); Kayakkatuyarykskij Creek 1600 m (8/
72), 1630 m (8/127); Kishte 440 m (Zolotukhin
11.VII.1988); Kokkul Lake 2300 m (33/10); Kokorya
Creek 2400 m (32/16); Kuba Creek, at mouth 500 m
(Makarov 3.VI.1972); Malyi Yaloman Creek, 4 km up-
stream 900 m (25/45); Tabozhok Peak 2250 m (31/151),
2750 m (31/75); Ulagan 1220 m (36/206); Ust-Sema
350 m (24/10), 400 m (24/129); Yailyu 440 m (0/639),
450 m (1/52), 500 m (0/996).
Entodon compressus C. Müll., Linnaea 18:
707. 1845. – E. challengeri (Par.) Card., Beih.
Bot. Centralbl. 17: 32. 1904. Figs. 2 & 3
Plants medium-sized, glossy, green. Stem prostrate,
complanately foliate, 3 cm long; central strand well-
developed; branches to 8 mm long, curved.
Pseudoparaphyllia broadly triangular. Axillary hairs
4-celled, 110 μm long, apical cell 45 × 4.5 μm. Stem
leaves 1.1-1.3 × 0.6-0.8 mm, ovate, obtuse to shortly
acute; margins entire or minutely serrulate near the
apex. Branch leaves 0.7-0.9 × 0.3-0.4 mm. Laminal
cells 55-75 × 6 (5/1) μm. Alar cells shorter in wide
area extending to the costa.
Autoicous. Sporophytes frequent. Inner pericha-
etial leaves greatly enlarged. Seta 0.5-0.7 cm. Cap-
sule ca. 2.0 mm long, ovoid, rugose.Operculum conic,
with long beak. Annulus of 2 rows of large cells,
deciduous. Exostome teeth straight when dry, gradu-
ally tapered, entire, densely low-papillose on both
surfaces; ventral trabeculae low. Endostome segments
with indistinct papillae. Spores 13 μm.
Distribution: This species comprises in general
an example of the classic East Asian – eastern North
American disjunction. In Asia it is widespread
throughout most of Japan (absent in Ryukyu) and
China (absent in Taiwan and Hunan), Korea, Mon-
golia, southern part of Russian Far East and South
Siberia westwards to Altai. Further to the west it
was collected once in Vologda Province in Cenrtal
Russia (LE!, H!) and was reported also from Cau-
casus (Düll, 1985). In Altai Entodon compressus is
28 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 3. Entodon compressus C. Müll. (from Maima 280 m Ignatov 35/43): 1 – perisome (450×); 2 – outer surface
of tooth near the base (5000×); 3 – outer surface of tooth in upper third (6500×); 4 – side view of tooth above the
middle, showing papillose inner surface and low ventral trabeculae (4600×).
3
2
1
4
strictly confined to flood valleys of large rivers where
Populus stands are developed. It grows mostly on
Populus trunks, or rarely on nearby rocks.
Specimens examined: Chulyshman River, at Bashkaus
mouth 470 m (Zolotukhin 23.IX.1989); Gorno-Altaisk 400
m (35/32); Kyrsai 450 m (Bardunov 6.VII.1966 IRK!;
Schischkin 7.IX.1931 LE!); Maima 280 m (35/18; 35/
21; 35/22; 35/43; 35/44; 35/59; 35/60).
Entodon schleicheri (Schimp.) Demeter,
Rev. Bryol. 12: 87. 1885. – Isothecium schle-
icheri Schimp. in Spruce, Musci Pyren. 71. 1847.
Figs. 4, 5 & 6
Plants robust, glossy and green. Stem prostrate,
complanately foliated, 8 cm long; central strand
present; branches to 8 mm long, straight. Pseudopara-
phyllia broadly triangular. Axillary hairs 4-celled,
180 μm long, apical cell 80 × 8 μm. Stem leaves 2.0-
2.5 × 0.8-1.1 mm, ovate, oblong-ovate, acute, narrowed
to the base; margins entire or minutely serrulate near
the apex. Branch leaves ca. 1.4 × 0.5 mm. Laminal
cells 100-120 × 6 (4/2) μm. Alar cells subquadrate,
forming a well-delimited triangular group, about 10-
15 along the margin.
Autoicous. Sporophytes frequent. Inner pericha-
etial leaves greatly enlarged. Seta 1.3 cm. Capsule ca.
2.5 mm long, long-cylindric. Annulus none or of 1
row on round cells with 40 μm in diameter. Opercu-
lum conic, with long beak. Exostome teeth straight
when dry, 0.35 mm long, gradually tapered, entire,
striolate below, densely papillose in the middle, smooth
above; ventral surface smooth. Endostome ca. 0.3 mm
long, segments smooth. Spores 13 μm.
In Altai this species is easy to recognize by
its large shiny complanate plants, usually with
numerous capsules. Several collections from (1)
strongly shaded and (2) very dry willow stands
in steppe areas were much thinner; without
sporophytes these specimens can be confused
with E. compressus, but the latter differs in the
smaller size of plants and arcuate branches.
29
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 4. Entodon schleicheri (Schimp.) Demeter (from Kobukhta 700 m Ignatov 0/247): 1 – habit; 2-3 –
perichaetial leaves; 4-6 – stem leaves; 7-9 – branch leaves; 10 – stem cross-section; 11 – middle laminal cells of stem
leaf; 12, 15 – pseudoparaphyllia; 13 – alar cells of stem leaf; 14 – upper laminal cells of stem leaf. Scale bars: 5 mm
– for 1; 1 mm – for 2-9; 100 μm – for 10-15.
5 mm
3
1
4
14 15
2
5
89
11
12
13
7
6
10
1 mm
100 μm
Krylov (1925) reported in Altai Entodon con-
cinnus (as E. orthocarpus), E. compressus and
E. cladorrhizans (Hedw.) C. Müll., the latter
name was probably applied to E. schleicheri. Bar-
dunov (1974) added E. schleicheri but did not
remove E. cladorrhizans, raising the total to there-
fore four species. However, we did not see any
specimens of true E. cladorrhizans in collections
from Altai, and therefore consider this species re-
ported erroneously.
Differences between E. schleicheri and E.
cladorrhizans.
In Europe both Entodon schleicheri and E.
cladorrhizans are rare mosses, thus providing
little opportunity for European bryologists of
the first half of XIX century to understand the
differences from North American plants of
Entodon cladorrhizans (holotype in G!).
30 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 5. Entodon schleischeri (Schimp.) Demeter from various regions (1, 3 – from Ural Mts., Bashkiria “Mosses of USSR, N
65” (MHA), Ignatova 13.IX.1990; 2, 7 – from Altai, Malyi Yaloman Creek 1050 m Ignatov 25/108; 4 – from Moscow Province,
Kunzevo, Zickendrath 27.IX.1989 (ex LE); 5-6 – from Altai, Chodro 880 m Zolotukhin 4.VII.1989): 1-2 – peristomes (1 – 200×;
2 – 270×); 3-4 – lower parts of exostome teeth, showing indistinct striolation (3-4 – 2100×); 5 – side view of exostome tooth in
upper part (2400×); 6-7 – lower parts of exostome teeth, showing distinct striolation (6 – 2500×; 7 – 3800×).
1 2
3 5
6
4
7
31
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 6. Entodon schleischeri (Schimp.) Demeter from various regions (1 – Switzerland, Baden, Alonz 1895 (ex LE),
2 – Russian Far East, Primorsij Territory, Lazovskij Reserve Czerdantseva 2.IV.1987 (ex VLA)): lower parts of teeth,
showing distinct striolation (1 – 5400×; 2 – 2700×).
1 2
Schimper was the first who found differ-
ences in annulus and exostome ornamentation
between European and North American plants.
He described Cylindrothecium schleicheri from
exsiccates of Spruce and later thoroughly ex-
plained the differences of these species from
American plants in Bryologia Europaea (1853).
According to Schimper C. schleicheri differs
in annulus absent (vs. present in C. cladorrhi-
zans) and exostome teeth striolate (vs. papil-
lose). However later he changed his mind and
synonymizied Cylindrothecium schleicheri with
C. cladorrhizans (Schimper, 1860). After that
some authors have followed the former idea of
Schimper, others – to the latter.
An important step toward understanding the
species was taken by Demeter (1884), who found
a specimen with annulus of 3 row of large cells
(1-2 rows of elongate cells with slit-like lu-
men) and non-striolate exostome, and described
it as Entodon transylvanicus Demeter. Later this
species was found to be identical with E. clad-
orrhizans (Limpricht, 1895-1904). Since that time
most European bryologists considered both E.
cladorrhizans and E. schleicheri present in Eu-
rope. However these taxa were treated in differ-
ent ways, so even in recent publications (Corley
& al., 1981) it has been noted that this problem
needs further study. In the North America E.
cladorrhizans is more common, while E. schle-
icheri have been revealed in few places only
recently (Buck & Crum, 1978). We have studied
several specimens of these species under both
light microscope and SEM.
Entodon cladorrhizans from North America
was found scarcely variable: all plants with a
well-separating annulus have irregularly retic-
ulate ornamentation. In E. schleicheri striola-
tion on the outer surface of exostome was found
rather variable. In most of specimens (from Far
East, Siberia, West Europe, North America) pa-
pillae are fused in rather solid cristae (Figs. 5
(6-7) & 6), resulting in a clear striolate pattern
well seen under light microscope. However in
some other specimens (Siberia, Urals, Central
Russia) rather low round and homogeneous
papillae form rather apparent rows, but solid
cristae are developed only in scattered places
or not at all (Figs. 5 (3-4)). Peristomes of the
latter form look, under the light microscope, as
indistinctly striolate to non-striolate. Neverthe-
less we attribute this form to E. schleicheri
because of (1) annulus absence or weak expres-
sion, and (2) gradual transitions in exostome
ornamentation. The form of E. schleicheri with
“indistinct-striolate” exostome has often been
considered as E. cladorrhizans s. str. (for ex-
ample in the Moscow Province of Russia). The
only specimen of E. cladorrhizans from Eur-
asia seen by us is the holotype of E. transyl-
vanicus (Fig. 7). All other non-American col-
lections in MHA, MW, LE and H studied be-
long to E. schleicheri. Several specimens were
seen from China, and all of them appear to be
32 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 7. Entodon cladorrhizans (Hedw.) C. Müll. from various regions (1-3 – Romania, isotype of E. transsylvanicus
Demeter (ex H); 4 – North America, New Mexico, Ikonberg 395 (ex NY)). 1 – peristome (360×); 2 – outer exostome
tooth in upper third (3600×); 3-4 – lower parts of teeth, showing reticulate ornamentation pattern (3 – 5300×; 4 –
4000×).
12
34
E. schleicheri. Since Hu (1983) mentioned for
China only one species of this pair (E. clador-
rhizans), we can conclude that only one species
occurs there, and it is E. schleicheri.
Distribution: Entodon schleicheri in the sense
mentioned above has scattered localities in several
countries of Central Europe (Austria, Germany,
France, Italy, Jugoslavia, Romania, Switzerland, Cze-
chia), Central Russia (Moscow Province), Urals and
Caucasus, and very common in mountains of South
Siberia, Russian Far East, Korea, and mainland Chi-
na (south to Yunnan). Also this species occurs in
western U. S. A. and in Mexico. In Altai is is very
common in more wet areas of Northern Altai, grow-
ing at lower elevations on fresh logs, rarely on trunks
of living trees, sometimes on rock outcrops (usually
on layer of humus), always in more or less exposed
habitats. In Central Altai this species occurs some-
times in rather xeric areas, but there only in valleys,
on Betula and Populus trunks.
Specimens examined: Anos 400 m (Anonymous
17.VIII.1909 LE!); Bele 500 m (0/67); Bolshoi Yaloman
Creek, in middle course 1500 m (Makarov & al.
15.VII.1972); Chemal 450 m (Schteinberg 8.IX.1931 LE!)
(29/74); Chemal Creek, 3 km upstream 450 m (34/64);
Chemal, 10 km upstream along Katun 450 m (34/55);
Chodro 880 m (Zolotukhin 4.VII.1989); Chulcha River,
33
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
100 μm
Fig. 8. Platygyrium repens (Brid.) B.S.G. (from Chiri Creek, 0.5 km upstream 450 m Ignatov 17/91): 1 – alar
cells of stem leaf; 2 – alar cells of branch leaf; 3 – habit; 4 – capsule; 5 – middle laminal cells of stem leaf; 6-8 –
stem leaves; 9-10 – perichaetial leaves; 11 – pseudoparaphyllia; 12 – upper lamina cells of stem leaf; 13 – upper
lamina cells of branch leaf; 14 – stem cross-section; 15-17 – branch leaves. Scale bars: 5 mm – for 3; 2 mm – for 4;
1 mm – for 9-10; 0.5 mm – for 6-8, 15-17; 100 mkm – for 1-2, 5, 11-14.
18
6
3
4
7
9
10
2
5 mm
2 mm
0.5 mm
15
13
14
12
11
17
16
in middle course 1000 m (9/134; 9/160), 1100 m (9/
122); Elekmonar Creek, 25 km upstream 1100 m (26/72);
Elekmonar Creek, 5 km upstream 700 m (26/39); Gorno-
Altaisk 400 m (35/13); Kairu Creek, 6 km upstream 900
m (15/152; 15/84); Kobukhta 700 m (0/247); Kyrsai
450 m (Schischkin 7.IX.1931 LE!); Maima 280 m (35/
19); Malyi Yaloman Creek, 5 km upstream 900 m (25/
118); Malyi Yaloman Creek, 7 km upstream 1050 m (25/
108); Yailyu 450 m (0/1000; 0/999), 470 m (0/998).
Platygyrium B. S. G.
The genus includes 3-5 species in temperate
regions of the Northern Hemisphere, one of which
has a wide distribution in the Holarctic. Platygy-
rium differs from Entodon in small plants, dio-
icous sexuality and brood branches in leaf axils.
Platygyrium is often considered to be close
to Pylaisiella. Their similarities include erect
capsules, peristome reduction and straight leaves.
However the peristome reduction in Pylaisiel-
la is confined only to imperfect cilia, while in
Platygytium the endostome has a low basal
membrane, with narrow, non-perforate segments.
Exostome teeth in Platygyrium are unusually
small for the Hypnaceae (0.15 mm, cf. 0.30 mm
in Pylaisiella polyantha), without typical pap-
illose to striolate ornamentation on the outer
surface and with smooth ventral trabeculae,
while in Pylaisiella the exostome teeth are pap-
illose, often with branched papillae, and with
incrassate papillose ventral trabeculae. Also,
Platygyrium and Pylaisiella differ in their
pattern of hygroscopic movement of the peris-
tome (in Platygyrium the teeth cover the mouth
when wet, while in Pylaisiella they are erect
when wet). So, there are no reasons to consider
these genera to be closely related.
Platygyrium repens (Brid.) B.S.G., Bryol.
Eur. 5: 98. t. 458. 1851. – Pterigynandrum repens
Brid., Muscol. Recent. Suppl. 1: 131. 1806.
Figs. 8 & 9
Plants small, dark-green in older parts, with light-
er and ±glossy young shoots. Stem to 2 cm long,
without central strand, ±regularly pinnate-branched,
branches 3-4 mm long. Pseudoparaphyllia lanceolate,
short. Axillary hairs 3-celled, 80 μm long, apical cell
40 × 11 μm. Rhizoids below leaf insertions or around
brood branches or in leaf axils where brood branch-
5
1 mm
50 μm
34 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 9. Platygyrium repens (Brid.) B.S.G. (from Moscow Province, “Mosses of USSR, N 30” (MHA) Ignatov
11.VI.1988): 1 – peristome (440×); 2 – side view of exostome tooth in upper third (3500×); 3 – outer surface of
exostome tooth in lower third (5200×); 4 – outer surface of exostome tooth in transition zone (5200×).
1
3
2
4
es could develop; some rhizoids granulose and pur-
plish. Stem leaves ± straight, 0.7-1.2 × 0.3-0.4 mm,
ovate, shortly acuminate; margins entire and recurved.
Branch leaves 0.4-0.6 × 0.2 mm. Laminal cells (35-)
60-90 × 8 (5/3) μm; in alar regions cells subquad-
rate, numerous.
Dioicous. Sporophytes rather frequent. Pericha-
etial leaves 1.4-1.5 × 0.4-0.5 mm, spreading; alar re-
gions wide and extending along the margin, forming
distinct borders. Seta 1.5 cm. Capsule ca. 1.5 mm
long, straight. Annulus of 2 rows of round cells. Oper-
culum conic-rostrate. Exostome teeth straight when
35
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
dry, 0.15 mm long, gradually tapered, entire; in the
lower 2/3 reticulate (proximally with more trans-
verse orientation of cristae, distally with more longi-
tudinal), in the upper part smooth to irregularly rug-
ose; dorsal trabeculae low; margins non-incrassate;
ventral trabeculae smooth. Endostome with very low
basal membrane, segments narrow and ca. 0.12 mm
long, cilia none. Spores 11-14 μm.
Well-developed extensive mats of Platygy-
rium are easy to recognize by rather small dull
dark-green plants with numerous brood branch-
es in leaf axils. However the young parts of
plants are ± shiny, Pylaisiella-like and usually
need microscopic study. Recurved leaf margins
are suggestive in this case. Fortunately P. repens
is often found with sporophytes and have setae
much longer than in Pylaisiella.
Distribution: Platygyrium repens is widespread in
the temperate and hemiboreal zones of Europe, North
Africa and North America. In Asia it occurs in South
Siberia and Russian Far East, Korea, Japan and in north-
ern and northwestern China. In Altai it occurs mostly
at elevations lower than 1200 m, avoiding true boreal
forests. Nearly all the collections were made from rotten
logs or from upper surfaces of inclined trunks.
Specimens examined: Adylda Creek 1100 m (34/79);
Anos 360 m (465; Vereshchagin 22.VI.1911 LE!); Bolshoi
Mianok Creek, at mouth 440 m (Zolotukhin 8.VIII.1988);
Chemal Creek, 3 km upstream 450 m (34/67; 34/68); Chiri
Creek, 0.5 km upstream 450 m (17/8; 17/91); Elekmonar
Creek, 5 km upstream 700 m (26/35); Kaitanak Creek 1200
m (Bondartzeva 1.VIII.1968 LE!); Korbu 440 m (Zolotukhin
20.X.1988); Kuba Creek, 5 km apart from Chemal 500 m
(Makarov 3.VI.1972); Ok-Porok Creek, at mouth 440 m (Zo-
lotukhina 30.III.1988); Shebalino 800 m (Seman 15.VII.1970
LE!); Shebalino deer farm 1000 m (Palamarchuk 23.VIII.1969
LE!); Yailyu 450 m (1/46), 480 m (0/625); between Ak-
Kem and Kochurla Creeks (Krylov 27.VII.1903 LE!).
Plagiotheciaceae (Broth.) Fleisch.
In the current literature the contents of this
family within the Holarctic is usually accept-
ed either for Plagiothecium + Isopterygium &
related genera (following Fleischer, 1912, etc.),
or for Plagiothecium only, leaving other gen-
era to Hypnaceae (Fleischer, 1923; Brotherus,
1925, etc). Recently Hedenäs (1987, 1989) found
that rhizoid topography and some other char-
acters define a group of genera of otherwise
unclear position – Myurella, Orthothecium,
Isopterygiopsis, Platydictya, and Herzogiella.
These genera appear close to each other, and
closer to Plagiothecium than to any part of the
Hypnaceae, so Hedenäs referred them to Pla-
giotheciaceae. This position is accepted in gen-
eral in this paper.
The Plagiotheciaceae are delimited by a num-
ber of synapomorphic characters, presented in at
least some species of the genera: (1) branching
rare, irregulary fasciculate; (2) branches fragile,
growing in the early stages at acute angle with
the stem; (3) lack of pseudoparaphyllia; (4) rhiz-
oid position in leaf axil or on the abaxial face of
leaf, or in indefinite position, but not below the
leaf insertion; (5) rhizoids papillose and purple;
(6) axillary hairs very thin; (7) stem and branch
leaves non-differentiated; (8) exostome teeth pale;
(9) outer surface of exostome above the transition
zone smooth or with rare minute papillae; (10)
gemmae in leaf axils or abaxial on leaf.
Hedenäs (1989) included Taxi phyllum
(and Isopterygium) in the Plagiotheciaceae.
However Taxiphyllum has (1) rhizoids below
the leaf insertion; (2) foliose pseudoparaphyl-
lia; (3) no gemmae; (4) more regularly pinnate
branching. We place this genus in Hypnaceae,
as did Brotherus (1925), Ireland (1986), Inoue
& Iwatsuki (1987), etc., although within the
Hypnaceae Taxiphyllum (and probably also
Isopterygium) has no obvious relatives.
Isopterygiopsis Iwats.
Iwatsuki (1970) described this genus for
only one species, I. muelleriana, but later add-
ed also I. pulchella (Iwatsuki, 1987), since it
was found closer to I. muelleriana, than to other
species of the large, heterogeneous Isopterygium.
Important differences from the latter include
(1) axillary papillose rhizoids; (2) lack of
pseudoparaphyllia; (3) epidermal layer of stem
composed of thin-walled cells. The genus in-
cludes 3 species, all present in Altai.
KEY TO THE ALTAIAN SPECIES OF
ISOPTERYGIOPSIS
1. Epidermal cells of stem as large as those of
the subtending layer; leaves indistinctly com-
planate, linear-lanceolate; plants autoicous,
often producing sporophytes
. . . . . . . . . . . . . . . . . . . . . . . . I. pulchella (p. 36)
1. Epidermal cells of stem markedly larger than
those of the subtending layer (see cross sec-
tion); plants dioicous, sporophytes never
found in Altai . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Plants small to medium-sized, distinctly
complanate . . . . . . I. muelleriana (p. 36)
2. Plants very small, with rather short leaf
cells . . . . . . . . . . . . . . . . I. alpicola (p. 38)
36 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 10. Isopterygiopsis muelleriana (Schimp.) Iwats. (from Kamga Creek, 14 km upstream 600 m Ignatov 0/111):
1-2 – habit; 3-6 – leaves; 7 – upper laminal cells; 8 – middle laminal cells; 9 – gemmae; 10 – stem cross-section;
11 – cells of leaf base. Scale bars: 2 mm – for 1-2; 0.5 mm – for 3-6; 100 μm – for 7-11.
1
2
34
6
5
10
8
11
9
7
2 mm
100 μm
0.5 mm
Isopterygiopsis muelleriana (Schimp.)
Iwats., J. Hattori Bot. Lab. 33: 379. 1970. – Pla-
giothecium muellerianum Schimp., Syn. Musc.
Eur. 584. 1860. Fig. 10
Plants small to medium-sized, intricate and green
or yellow-green, distinctly complanate. Stem pros-
trate, ca. 2.5 cm long, with hyalodermis and central
strand, irregularly loose branched. Axillary hairs 3-
celled, 190 μm long, apical cell to 95 × 7-9 μm. Leaves
0.7-0.9 × 0.2-0.3 mm, oblong-ovate, entire, densely ar-
ranged, usually overlapping. Laminal cells 40-60 ×
3.5 μm. Alar cells not differentiated. Gemmae in leaf
axils 4-5-celled, uniseriate, 55-75(-85) × 9-11 μm.
Dioicous. No sporophytes have been seen in Alta-
ian plants (or in neighboring regions).
Isopterygiopsis muelleriana is easy to know
by small to medium-sized plants with a dis-
tinctly complanate habit, growing mostly on wet
rocks. Very characteristic are some parts of stem
where leaves are much smaller; such zones are
usually on shoot ends, but sometimes they grow
further into normally foliated shoots.
Distribution: Isopterygiopsis muelleriana has a
scattered distribution throughout the Holarctic, in
more southern mountains and oceanic regions. In
South Siberia and Russian Far East it is known
from nearly all the places where careful studies were
undertaken; the species is known from South and
Central Europe, Caucasus, Himalayas, Sichuan, north-
eastern China, and Japan. However it was not found
in Mongolia and Xinjiang, probably because of gen-
erally xeric conditions. In Altai it has been found in
the forest- and subalpine belts, typically on moist or
wet sides of rock outcrops (usually on ±basic schists).
Specimens examined: Bayas Lake 1750 m (0/240);
Bolshoi Shaltan Creek 580 m (0/1241); Kamga Bay 450
m (Zolotukhin 11.VII.1988); Kamga Creek, 14 km up-
stream 600 m (0/111), 630 m (0/1251); Kayakkatuyaryk-
skij Creek 1760 m (8/14), 1800 m (8/228), 1900 m (8/
315), 2000 m (3/41), 2050 m (7/39); Kobiguayuk Creek
2180 m (0/1237); Maloye Istyube Creek, 1 km upstream
600 m (Zolotukhin 19.V.1977); Srednij Shaltan Creek
730 m (0/1243); Yailyu 450 m (0/1252; 1/17).
Isopterygiopsis pulchella (Hedw.) Iwats.,
J. Hattori Bot. Lab. 63: 450. 1987. – Leskea
pulchella Hedw., Sp. Musc. 220. t. 55: f. 7-12.
1801. Figs. 11 & 12
Plants small, green to yellow-green. Stem 2-2.5
cm long, irregularly branched; epidermal cells small,
but rather thin-walled; central strand present. Axil-
lary hairs 2-celled, 65 μm long, apical cell 57 × 6 μ m.
Leaves erect-spreading to somewhat falcate (in more
xeric conditions), (0.5-)0.8-1.0 × 0.2 mm, lanceolate
to ovate-lanceolate. Laminal cells 85-130 × 4 μm. Basal
cells shorter, not differentiated in alar regions. Gem-
mae in leaf axils, 3-4-celled.
Autoicous. Sporophytes frequent. Inner pericha-
etial leaves to 0.5 × 0.1 mm, smooth. Seta 0.8-1.2 cm.
37
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 11. Isopterygiopsis pulchella (Hedw.) Iwats. (from Tabozhok Peak 2350 m Ignatov 31/99): 1 – habit; 2-
3 – leaves; 4-6 – perichaetial leaves; 7 – middle laminal cells; 8 – upper laminal cells; 9 – stem portion; 10 – stem
cross-section; 11 – cells of leaf base. Scale bars: 5 mm – for 1; 2 mm – for 9; 1 mm – for 4-6; 0.5 mm – for 2-
3; 100 μm – for 7-8, 10-11.
10
5
8
11
9
6
1
2
3
4
7
5 mm
2 mm
0.5 mm
1 mm
100 μm
Capsule ca. 1.0 mm long, straight to rarely somewhat
curved. Annulus of 2 rows of cells, deciduous. Opercu-
lum conic. Exostome teeth ca. 0.35 mm long, gradually
tapered, entire, reflexed when dry; in the lower part
striolate, with cristae wavy and somewhat papillose;
in the middle cristae becoming oblique to longitudinal
and with overlying papillae, or sometimes forming a
reticulate pattern (with longitudinally elongated
loops); in the upper part with rare low irregularly
arranged papillae, low dorsal trabeculae, non-incras-
sate margin, and smooth ventral surface and trabecu-
lae. Endostome 0.40 mm long, basal membrane 0.10
mm high, segments narrow and non-perforate; cilia
rather short and nodose to reduced. Spores 10-15 μm.
Isopterygiopsis pulchella can be recognized
in the field by numerous pale capsules, similar
to those of Plagiothecium laetum, and subjula-
ceous habit.
Distribution: Isopterygiopsis pulchella is a moss
of cool climate of both hemispheres. It is widespread
in the Arctic (including Svalbard, Franz-Josef Land,
northern Greenland, etc.) and northern boreal for-
ests and also high mountains throughout the Hol-
arctic. It is common in Mongolia, but in China was
reported only from the northern provinces,
Heilongjiang and Xinjiang. In Altai I. pulchella is
very common in the taiga belt at ca. 1500-2000 m,
where it grows on rotten logs and on soil at up-
turned roots of fallen trees. It is not rare locally in
the alpine zone, especially in more xeric areas of south-
eastern Altai, where it grows among rocks as well as
on soil in tundras.
Specimens examined: Bashkaus River, upper course
2250 m (36/380a); Bayas Lake 1750 m (0/659), 2100 m
(0/1036); Bogoyash Creek, upper course 2400 m (36/
334a), 2550 m (36/246); Chainary Creek, upper course
1800 m (34/196); Chiri Creek, 0.5 km upstream 450 m
(17/111); Chulcha River, in middle course 850 m (Zolo-
tukhin 10.IX.1989); Kairu Creek, 8 km upstream 1000 m
(15/101; 15/48; 15/90); Karakol Lakes 1900 m (26/
122; 28/71); Kayakkatuyarykskij Creek 1650 m (8/131),
1900 m (8/320; 8/337), 1920 m (3/159; 3/219; 3/
253), 1940 m (3/243), 2050 m (3/4; 7/147); Kobigu-
ayuk Creek 2030 m (0/1037), 2200 m (0/1038; 0/1707),
2500 m (0/301); Kokkul Lake 2300 m (33/13); Kokorya
Creek 2150 m (32/19), 2400 m (32/18); Kukol 1780 m
(0/1035), 1800 m (0/1034; 0/1039), 1850 m (0/660);
Malaya Kokorya Creek 2450 m (36/287); Saluru Creek,
upper course 2250 m (36/293); Sondrukat Creek, upper
course 2000 m (Zolotukhin 13.VI.1988); Tabozhok Creek,
12 km upstream 2200 m (30/46); Tabozhok Peak 2150 m
(31/105), 2300 m (31/103; 31/98), 2350 m (31/100;
31/99), 2550 m (31/97), 2750 m (31/102); Tokpak Creek,
in middle course 2800 m (36/389a); Yailyu 450 m (1/
9); Yakhansoru Lake 2000 m (Zolotukhin 26.VI.1990).
38 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1
3
2
4
Fig. 12. Isopterygiopsis pulchella (Hedw.) Iwats. (from Saluru Creek 2250 m Ignatov 36/293): 1 – peristome
(350×); 2 – inner surface of exostome tooth in upper part, showing smooth inner trabecula (4800×); 3 – outer
surface of exostome tooth above the transition zone (3000×); 4 – outer surface of exostome tooth below the transition
zone (4800×).
Isopterygiopsis alpicola (Lindb. et H. Ar-
nell) Hedenäs, J. Bryol. 15: 495. 1988. – Stere-
odon alpicola Lindb. et H. Arnell, Kongl. Sv.
Vet. Acad. Handl. 23(10): 153. 1890. Fig. 13
Holotype: Siberia, Jenisei, Uskij Mys, 61°30'
N, in schists 5.VII.1876 H. W. Arnell (S!).
Plants small, in rather dense green opaque tufts.
Stem 1.5 mm long, loosely foliate, with strongly en-
larged epidermal cells, with central strand, 15 mm
long, very rarely branched. Axillary hairs 2-celled,
50 μm long, apical cell 45 × 4 μm. Leaves somewhat
falcate, 0.3-0.35 × 0.1-0.15 mm, ovate, shortly acumi-
nate; margins serrulate throughout. Laminal cells 25-
40(-55) × 6-8 (5-6/1-2) μm, often irregular in shape.
Alar cells shorter, forming a small indistinct group.
Dioicous? Sporophytes unknown. Gemmae in leaf
axils 3-celled, ca. 45 × 9 μm.
Isopterygiopsis alpicola is a small plants, su-
perficially somewhat similar to small forms of I.
pulchella, but differs in dioicous sexual condition,
serrulate leaves, shorter laminal cells and large
epidermal cells. The differences from Platydictya
jungermannioides include the presence of central
strand (ill-developed in thinner shoots), large
epidermal cells of the stem, and differences in
gemma shape (cf. Figs. 13 & 14) – in Isopterygi-
opsis gemmae are composed of a row of 3-4 elon-
gate cells of the same width, while in Platydictya
the cells are shorter, and gemmae have an obcu-
neate shape due to the wider distal end of the
distal cell, which is often slightly lobed.
39
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 13. Isopterygiop-
sis alpicola (Lindb. et H.
Arnell) Hedenäs (from
Kayakkatuyarykskij Creek
1900 m Ignatov 8/337):
1 – habit; 2 – gemmae; 3
– stem cross-section; 4-5
– stem leaves. Scale bars:
2 mm – for 1; 100 μm –
for 2-5.
1
3
5
2
2 mm
Distribution: This is a very rare moss world-
wide. We have seen three specimens from Siberia ((1)
type; (2) one collection from Altai; (3) Middle course
of Yenisej River, 63°20’N-91°9'E, Szerbina 139, MW);
one collection from Amurskaya Province of the Rus-
sian Far East (53°52'N-127°19'E, 350 m alt., L. I.
Abramova, s. n., MW!); and several collections from
Sweden and Finland (in S!).
Specimens examined: Kayakkatuyarykskij Creek 1900
m (8/337).
Platydictya Berk.
For a long time Amblystegiella, nom. illeg.
and later Platydictya have been considered in
a wide sence, including small mosses without
central strand and with costa faint or absent.
However Hedenäs (1987) showed that P.
jungermannioides, the type of the genus, is dis-
tinct from other widespread Holarctic species,
P. subtilis (Hedw.) Crum and P. confervoid-
es (Brid.) Crum. As to the latter two species,
Söderström & al. (1992) suggested to place them
in Serpoleskea (Limpr.) Warnst., a genus typ-
ified by Leskea subtilis Hedw. (= Serpoleskea
subtilis (Hedw.) Loeske). It needs a further
study to understand if other Chinese and North
American species referred to Platydictya be-
long to this genus or to Serpoleskea.
Among the pleurocarps with axillary rhiz-
oids Platydictya is peculiar in minute size of
plants and non-differentiated epidermal cells.
100 μm
Fig. 14. Platydictya jungermannioides
(Brid.) Crum (from Kukol 1800 m Ig-
natov 0/455): 1 – habit; 2 – portion
of rhizoid; 3 – axillary gemmae; 4 –
stem cross-section; 5-6 – leaves. Scale
bars: 2 mm – for 1; 100 μm – for 2-6.
156
3
4
2
2 mm
100 μm
4
40 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Platydictya jungermannioides (Brid.)
Crum, Michigan Bot. 3: 60. 1964. – Hypnum
jungermannioides Brid., Muscol. Recent. Suppl.
2: 255. 1812. Fig. 14
Plants very small, in loose green to yellow-
green dull tufts or growing among other bryo-
phytes. Stem ca. 1 cm long, loosely foliate, irregu-
larly branched. Axillary hairs 2-celled, 35 μm long,
apical cell 30 × 4.0 μm. Leaves straight, spread-
ing, 0.2-0.3 × 0.06-0.09 mm, lanceolate to ovate-
lanceolate, acuminate; margins serrulate, especial-
ly near the base. Laminal cells rhombic, 15-25 × 6-
7 (5-6/1) μm. Alar cells in a small inconspicu-
ous group. Gemmae in leaf axils 3-celled, 27-35
μm long, 7-8 μm wide in the middle, 12-17 μm wide
in their distal clavate end.
Dioicous. Sporophytes not seen in Altai. Micro-
graphs of the exostome was published by Ignatov &
Ochyra (1995): exostome teeth long, abruptly nar-
rowed in the transition zone, entire above; in the low-
er part striolate, with ±perfect, waved cristae; in the
middle densely papillose; in the upper part with rare
small papillae and prominent dorsal trabeculae.
Distribution: Platydictya jungermannioides
occurs across the Holarctic in the Arctic and boreal
zones, and in high mountains (in Central Asia, Cau-
casus, Yunnan, etc.). In Altai this species is infre-
quent at middle and upper elevations, on mesic soil
banks and among rocks.
Specimens examined: Bogoyash Creek, upper course 2350
m (36/264); Kaitanak Creek 1500 m (A.Krylov & Reczan
10.VII.1968 LE!); Kukol 1800 m (0/455); Tabozhok Peak
2250 m (31/163), 2750 m (31/164; 31/169; 31/317).
Myurella B. S. G.
The genus includes 4 species, growing in regions
with cool climate, and is characterized by: (1) plants
small, very fragile, julaceous; (2) leaves nearly round,
strongly concave; (3) laminal cells short, papillose at
back; (4) plants dioicous, with very rare sporophyte
production; (5) capsule erect. Besides the normally
developed imbricate shoots, all species produce stolonif-
erous shoots with rather remote leaves, which are dif-
ferent from “stem” leaves in (a) ±appressed to the
stem in their base and with spreading acumen; (b)
±ovate to lanceolate and longer-acuminate; (c) lam-
inal cells longer. No sporophytes were found in Alta-
ian plants of Myurella; micrographs of exostome of
all three species known in Altai were published by
Ignatov & Ochyra (1995).
This genus was placed for a long time in
different families (Thuidiaceae, Theliaceae, Pteri-
gynandraceae). Hedenäs (1987) suggested its
relation to the Plagiotheciaceae, and Ignatov &
Ochyra (1995) provided additional evidence
for a relationship to Platydictya.
KEY TO THE ALTAIAN SPECIES OF MYURELLA
1. Leaves rounded-obtuse, closely vermicular-
imbricate . . . . . . . . . . . . . M. julacea (p. 41)
1. Leaves acuminate, freely imbricate to spread-
ing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Leaf margins serrulate; laminal cells with
a low papilla in the upper corner on the
dorsal surface . . . . M. tenerrima (p. 42)
Fig. 15. Myurella julacea (Schwaegr.) B.S.G. (from Kukol 1750 m Ignatov 0/1048): 1 – habit; 2-5 – leaves; 6 – upper
laminal cells; 7 – stem cross-section; 8 – alar cells. Scale bars: 1 mm – for 1; 0.2 mm – for 2-5; 100 μm – for 6-8.
5
8
6
1
2
3
4
7
1 mm 0.2 mm
100 μm
41
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 16. Myurella sibirica (C. Muell.) Reim. (from Kairu Creek, 8 km upstream 1000 m Ignatov 15/14): 1 – habit;
2-4 – leaves; 5 upper laminal cells; 6 – middle laminal cells; 7 – stem cross-section; 8 – alar cells; 9-10 – perichaetial
leaves. Scale bars: 1 mm – for 1; 0.2 mm – for 2-4, 9-10; 100 μm – for 5-8.
1
2
58
6
34
7
9
10
1 mm 0.2 mm
100 μm
2. Leaf margins ciliate-dentate; laminal cells
with a high central papilla on the dorsal
surface . . . . . . . . . . . . . M. sibirica (p. 41)
Myurella julacea (Schwaegr.) B.S.G., Bry-
ol. Eur. 6: 41. 1853. – Leskea julacea Schwae-
gr. in Schultes, Reise Glockner 2: 363. 1804.
Fig. 15
Plants small, in pure whitish-green to glaucous
tufts or often growing among other mosses. Stem
vermicular-imbricate, usually 1 cm long, rarely ir-
regularly branched. Axillary hairs 4-celled, 100-110
μm long, apical cell 35 × 4 μm. Leaves 0.3-0.4 × 0.3-
0.4 mm, broadly-ovate to orbicular, obtuse, strongly
narrowed toward insertion, very concave, ecostate;
margin serrulate. Laminal cells 10-17 × 7 (5/2) μm,
with an obtuse papilla in the upper corner; margin-
al cells with thinner walls, forming a distinct border
2 cells wide; alar cells not differentiated. Dioicous.
No sporophytes have been found in Altai or neigh-
boring regions. Perichaetia numerous.
Distribution: Myurella julacea is widespread
throughout cool regions of the Holarctic, and also
in the Southern Hemisphere in Argentina and Ant-
arctic islands (for worldwide map see Ochyra &
Szmajda, 1990). It is common in the Arctic and moun-
tain regions, but absent or very rare in lowlands where
there are no rock outcrops. This is a very common
species throughout the Altai, in both wet and dry
habitats, mostly on rocks.
Specimens examined: Bayas Lake 1850 m (0/1050);
Berekhtuyaryk 1600 m (0/1134); Bogoyash Creek, upper
course 2350 m (36/87), 2400 m (36/51), 2450 m (36/82);
Chulcha River, in middle course 1100 m (9/84); Kairu Creek,
8 km upstream 1000 m (15/28); Kairu-Bazhi Peak 1700 m
(13/90), 2100 m (13/164; 13/53; 13/72; 13/87), 2300 m
(13/137); Karagai 500 m (Zolotukhin 10.VIII.1988); Kar-
akol Lakes 1900 m (26/127); Kayakkatuyarykskij Creek
2000 m (8/155), 2150 m (7/119); Kobiguayuk Creek 2100
m (0/1051), 2300 m (0/320); Kokorya Creek 2400 m (32/
10; 32/9); Kukol 1750 m (0/1048; 0/1049), 1800 m (0/
1047; 0/1128; 0/667); Maima 350 m (35/5); Tabozhok
Peak 2150 m (31/32), 2250 m (31/30), 2300 m (31/28;
31/33), 2350 m (31/25), 2550 m (31/24; 31/27), 2700 m
(31/31), 2750 m (31/29), 2800 m (31/26).
Myurella sibirica (C. Müll.) Reim., Hedwi-
gia 76: 292. 1937. – Hypnum sibiricum C. Müll.,
Syn. Musc. Frond. 2: 418. 1851. Fig.16
Plants small, light-green. Stem julaceous, to 1 cm
long, rarely irregularly branched. Leaves 0.3-0.4 ×
0.3-0.6 mm, broadly-ovate to orbicular, shortly acumi-
nate, strongly narrowed toward insertion, strongly
concave, ecostate; margins ciliato-dentate. Laminal
cells 12-22 × 8-10 (6-8/1.5-2) μm, with one high cen-
tral papilla. Dioicous. No sporophytes have been found
in Altai or neighboring regions.
Distribution: Myurella sibirica is widespread
in mountain regions of South Siberia, China, Japan,
and Himalayas, and less widespread in Europe. Com-
pared to M. tenerrima this is a more southern spe-
cies, very rare in the Arctic and occurring in lower
belts of mountains. In Altai it grows on wet soil and
rocks, mostly at lower elevations.
Specimens examined: Bolshoi Shaltan Creek 550 m
(0/1129; 0/1130); Derbogach 450 m (0/17; 0/26);
Ezhon Cape 440 m (Schischkin 10.IX.1931 LE!); Kairu
Creek, 8 km upstream 1000 m (15/125; 15/14; 15/82);
Kairu-Bazhi Peak 2100 m (13/71); Kamga Creek, 12 km
upstream 550 m (0/106); Kamga Creek, upper course 830
m (Zolotukhin 7.VI.1991); Karagai 450 m (Zolotukhin
11.VIII.1988), 500 m (Zolotukhin 10.VIII.1988); Kishte
42 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
440 m (Zolotukhin 11.VII.1988); Korbu 440 m (Zolo-
tukhin 15.VII.1988); Kukol 1800 m (0/1131), 1900 m
(0/1132); Maloye Istyube Creek, 1 km upstream 600 m
(Zolotukhin 19.V.1977); Ok-Porok Creek, 1.5 km upstream
470 m (2/47).
Myurella tenerrima (Brid.) Lindb., Musci
Scand. 37. 1879. – Pterigynandrum tenerrimum
Brid., Muscol. Recent. Suppl. 4: 132. 1819[1818].
Fig. 17
Plants small, pale-green. Stem julaceous, to 1 cm
long, rarely irregularly branched. Leaves 0.3-0.6 ×
0.3-0.4 mm, broadly-ovate to orbicular, abruptly short-
ly acuminate, strongly narrowed toward insertion,
strongly concave, ecostate; margins serrulate. Lami-
nal cells 11-18(-23) × 7-9 (5-7/2) μm, rhombic, with
a papilla in the upper corner. Alar cells not differen-
tiated. Dioicous. No sporophytes have been found in
Altai or neighboring regions.
Usually Myurella sibirica and M. tenerrima are
quite distinct, and occur in different zones. However,
sometimes poorly developed forms of Myurella si-
birica do not have especially long teeth on the leaf
margins and may be confused with M. tenerrima.
The most useful character to identify M. sibirica seems
be the central position of papillae on laminal cells.
Distribution: Myurella tenerrima is principally
an Arctic species, occurring in the alpine belt of some
southern mountains (for worldwide map see Ochyra
& Szmajda, 1990). In Altai it is rare in the subalpine
and alpine zones, on rocks and soil.
Specimens examined: Bayas Lake 1750 m (0/1133);
Kairu Creek, upper course 1700 m (13/195; 13/91).
Orthothecium B. S. G.
This genus includes about 10 species, grow-
ing principally in cold and cool climates in
the Northern Hemisphere.
Orthothecium was placed for a long time in
the Entodontaceae or Hypnaceae. However, the
peristome of Orthothecium is ±perfect, with typ-
ical hypnoid teeth and endostome with high
basal membrane. Therefore it has nothing to do
with the former family. The combination of char-
acters of species with axillary rhizoids suggests
a closer relation to Plagiotheciaceae (Hedenäs,
1987) than to Hypnaceae. Within the group of
genera with axillary rhizoids Orthothecium is
peculiar in the stem with thick-walled epider-
mal cells; leaves ±longly acuminate, truncate at
base; laminal cells narrowly linear; often bronze
to red color of plants. There is a general habit
and size similarity between small-sized species
of this genus, like O. strictum, and Isopterygi-
opsis. Large -sized species (O. rufescens (Brid.)
B. S. G. and O. chryseum) have many apomor-
phic characters (robust plants, rigid and strongly
plicate leaves). Besides axillary, rhizoides on abax-
ial side of leaf base occur in O. rufescens.
Fig. 17. Myurella tenerrima (Brid.) Lindb. (from Bayas Lake 1750 m Ignatov 0/1133): 1 – habit; 2 – middle
laminal cells at leaf margin; 3-5 – leaves; 6 – upper laminal cells; 7 – middle laminal cells; 8 – alar cells. Scale bars:
1 mm – for 1; 0.2 mm – for 3-5; 100 μm – for 2, 6-8.
4
1
7
35
8
6
100 μm
2
0.2 mm
1 mm
43
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
100 μm
Fig. 18. Orthothecium chryseum (Schwaegr.) B.S.G. (from Tabozhok Peak 2700 m Ignatov 31/95): 1 – habit; 2 –
branch initial with proximal branch leaves; 3 – upper laminal cells; 4 – middle laminal cells; 5 – alar cells; 6-8 – leaves;
9 – proximal branch leaf; 10 – stem cross-section. Scale bars: 1 cm – for 1; 1 mm – for 6-8; 100 μm – for 2, 3-5, 9-10.
1
5
8
6
34
79
2
10
1 cm
1 mm
Orthothecium has been considered to have
pseudoparaphyllia (Hedenäs, 1987); however
this character seems open to interpretation. The
youngest branch initials are covered by incised
lanceolate to triangular leaf-like structures. Af-
terward, the proximal branch somewhat elon-
gates, so all these structures appear above the
leafless narrowed zone with no foliose struc-
tures below the ring of small cells at branch
base (Fig. 18–2). This situation seems more sim-
ilar to those observed in Isopterygiopsis and
Myurella. So we interpret these leaf-like struc-
tures as juvenile leaves and pseudoparaphyllia
therefore are lacking.
KEY TO THE ALTAIAN SPECIES OF
ORTHOTHECIUM
1. Robust plants, with strongly plicate leaves
. . . . . . . . . . . . . . . . . . . . . . . O. chryseum (p. 43)
1. Rather small plants, with non-plicate leaves
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.
2. Leaves long-acuminate, with plane mar-
gins . . . . . . . . . . . . O. intricatum (p. 45)
2. Leaves short-acuminate, with recurved
margins . . . . . . . . . . . O. strictum (p. 45)
Orthothecium chryseum (Schwaegr.)
B.S.G., Bryol. Eur. 5: 107. 1851. – Hypnum chry-
seum Schwaegr. in Schultes, Reise Glockner 2:
364. 1804. Fig. 18
Plants rather robust, in ±soft reddish-golden shiny
tufts. Stem ascending to erect, 6 cm long, without
hyalodermis, with central strand, rarely branched;
young branches departing from the stem at acute
angle and often then ±parallel to the stem. Axillary
hairs 3-4-celled, 77-100 μm long, apical cell 30-50 × 4
μm. Leaves straight, ±appressed, 2.8 × 1.0 mm, broad-
ly lanceolate, deeply plicate; margins entire and in-
curved nearly throughout. Laminal cells 60-90 × 5-7
(1.5-3/3-5) μm, with very thick walls. Basal cells
usually colored, shorter; alar groups of subquadrate
cells small and indistinct. Dioicous. Sporophytes
unknown in Altai or neighboring regions.
According to numerous literature records, O.
rufescens is rather common in the Asian Arctic
(Afonina & Czernyadjeva, 1995) and it has been
reported by Bardunov (1974) in the Sayan and
Altai mountains in South Siberia. However we
found all so-named collections from Asian
Russia to be in fact O. chryseum. The latter
species is very polymorphic in Altai and repre-
sented mostly by forms with long, gradually
tapering leaves. However (1) there are numer-
44 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Table 1. Differences between Orthothecium chryseum and O. rufescens
O. chryseum O. rufescens
plant color bronze- or golden-reddish brownish- or purplish-red
leaves when dry appressed on most parts of the stem, spreading along practically all
spreading in uppermost part of stem the stem length
leaf margins at base plane, although sometimes masked incurved
by submarginal plicae
alar cells ±differentiated non differentiated
median laminal cells mostly shorter than 100 μm long mostly 110-150 μm long
ous transitions to the typical, shortly pointed
form which could be found in any extensive
collections; (2) leaves of proximal shoot are
constantly short-acuminate, though distal leaves
became often long-acuminate. So though the leaf
shape is widely used in keys, it has a restricted
taxonomic value. The descriptions of plant col-
or, as golden in O. chryseum, reddish in O. rufe-
scens, are also misleading, since O. chryseum in
sunny places is usually red. The more stable char-
acters are summarized in table 1.
We have seen specimens of O. rufescens from
Central Europe, Scandinavia and Japan.
Distribution: Orthothecium chryseum has a wide
distribution in the Arctic (including northernmost
very severe regions) and high mountains of Europe,
Siberia, Central Asia and North America. In Altai it
grows in wet open tundras in southeastern Altai,
100 μm
Fig. 19. Orthothecium intricatum (Hartm.) B.S.G. (from Karakol Lakes 1950 m Ignatov 28/80): 1 – habit; 2-5 –
leaves; 6 – upper laminal cells; 7 – middle laminal cells; 8, 10-11 – proximal branch leaves; 9 – stem cross-section;
12 – alar cells. Scale bars: 5 mm – for 1; 1 mm – for 2-5; 100 μm – for 6-12.
10
1
5
8
6
3
4
7
911
12
2
5 mm
1 mm
45
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
100 μm
Fig. 20. Orthothecium strictum Lor. (from Kukol 1800 m Ignatov 0/1020): 1 – habit; 2-4 – leaves; 5 – upper
laminal cells; 6 – middle laminal cells; 7 – stem cross-section; 8-9 – alar cells. Scale bars: 5 mm – for 1; 0.5 mm –
for 2-4; 100 μm – for 5-9.
1
5
8
6
34
7
9
2
5 mm 0.5 mm
rarer in alpine regions of other parts of Altai. The
second type of habitat of this species is the moist
“dripping” cliffs, where it grows at all elevations.
Specimens examined: Bogoyash Creek, upper course 2350 m
(0/2047; 36/197; 36/218), 2400 m (36/254); Bolshoi Shaltan
Creek 540 m (Zolotukhin 7.VI.1989), 560 m (0/1019; 0/641);
Bolshoye Istyube Creek 470 m (Zolotukhin 5.VII.1988; 18/
119); Kairu-Bazhi Peak 1700 m (13/193), 2100 m (13/39);
Karakol Lakes 1950 m (28/99); Kukol 1850 m (0/1018);
Tabozhok Peak 2350 m (31/91), 2450 m (31/88), 2700 m (31/
94; 31/95), 2750 m (31/89; 31/92), 2800 m (31/90; 31/93).
Orthothecium intricatum (Hartm.) B.S.G.,
Bryol. Eur. 5: 108. 457. 1851. – Leskea intrica-
ta Hartm., Handb. Skand. Fl., ed. 5, 336. 1849.
Fig. 19
Plants rather small in ±glossy, green to reddish-
golden tufts. Stem 1.0-1.5 cm long in upper leafy
part, with central strand, rarely branched. Axillary
hairs 3-celled, 70 μm long, apical cell 30 × 4 μm. Leaves
erect, 1.5-1.8 × 0.3-0.4 mm, lanceolate, longly acumi-
nate; margins entire, recurved at places. Laminal cells
50-70 × 6 (4/2) μm. Alar cells not differentiated.
Dioicous. Sporophytes are unknown in Altai or neigh-
boring regions.
Distribution: Orthothecium intricatum occurs
mostly in mountains of boreal to temperate zone,
with only few localities in the Arctic. It is known in
mountains of Europe and northern Africa, Caucasus,
Crimea, Middle Asia, Tibet, Himalayas, and Japan. In
Altai it was found on moist rocks in the forest to
subalpine zones.
Specimens examined: Derbogach 440 m (0/1021);
Kamga Creek, 10 km upstream 560 m (0/1022); Karakol
Lakes 1950 m (28/80).
Orthothecium strictum Lor., Moosstudien,
122. f. 5d. 1864. Fig. 20
Plants very variable, from medium-sized exten-
sive mats with ±prostrate stems ca. 2.5 cm long to
small dense tufts with stems less than 1 cm, ±glossy,
green to reddish-golden. Stem rarely branched, with
central strand. Axillary hairs 3-4-celled, 80-100 μm
long, apical cell 30-40 × 4 μm. Leaves erect to tightly
appressed (in smaller forms), 1.0-1.3 × 0.3 mm, lan-
ceolate, shortly acuminate; margins entire, recurved
at places or nearly throughout (in smaller forms).
Laminal cells (40-)50-85 × 6 (4/2) μm. Alar cells
not differentiated. Dioicous. Sporophytes unknown
in Altai or neighboring regions.
Orthothecium strictum is polymorphic in
Altai. Alpine forms are very small and have
densely imbricate leaves, while plants from more
sheltered places and from lower altitudes are
much larger and have erect-spreading leaves.
This variability is easy to interpret as an envi-
ronmental response.
Distribution: This is mostly an Arctic species,
rarely found in mountains of northern Europe, Urals
46 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
100 μm
Fig. 21. Plagiothecium cavifolium (Brid.) Iwats. (“cavifolium-phenotype”: 1-2, 7-13 – from Yailyu 460 m Ignatov
0/1242; “succilentum-phenotype”: 3-6 – from Kobukhta 700 m Ignatov 0/248): 1-3 – habit; 4-8 – leaves; 9 – middle
laminal cells; 10-11 – upper laminal cells; 12 – alar cells; 13 – stem cross-section. Scale bars: 5 mm – for 1-3; 1 mm
– for 4-8; 100 μm – for 9-13.
13
1
5
8
6
3
4
7
910
11
12
2
5 mm
1 mm
(southward to about 60°N), and Siberia. In Altai
Orthothecium strictum grows on soil and rocks,
mostly above the tree line, with only one locality in
the upper forest zone.
Specimens examined: Ak-Turu Creek, upper course 2800
m (Bardunov 19.VII.1966 IRK!); Bogoyash Creek, upper course
2450 m (36/75), 2500 m (36/80); Kukol 1800 m (0/1020);
Tabozhok Peak 2700 m (31/319), 2750 m (31/183; 31/96);
Tokpak Creek, in middle course 2050 m (36/42).
Plagiothecium B. S. G.
Plagiothecium includes about 30 species, dis-
tributed in cold, boreal and temperate zones of
both hemispheres. It is a well-delimited genus,
characterized by: (1) stem with thin-walled epi-
dermal cells; (2) tendency to complanate foliage
and asymmetric leaves; (3) decurrent leaves; (4)
rhizoids mostly abaxial on leaves, often on costa.
The segregation from Plagiothecium is pos-
sible for P. latebricola, into the monotypic ge-
nus Philoscia Berk., Handb. Brit. Moss. 146. 1863.
This species has an isolated position due to (1)
reduction of endostome, which lacks cilia, and
has low basal membrane; (2) exostome teeth
with longitudinal cristae in the middle and lon-
gitudinal striolation above the transition zone;
ventral trabeculae smooth; (3) spore release in
winter, not in summer, as in all other species at
least in our territory; (4) gemmae on leaf tips
and in leaf axils, but not abaxial on leaves.
47
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1 2
3 4 5
Fig. 22. Plagiothecium cavifolium (Brid.) Iwats. (from Ukraine, Lvov Province, Ignatov 14.IX.1991, MHA): 1 –
peristome (220×); 2 – side view of exostome tooth in upper part, showing papillose inner trabeculae (3800×); 3 – outer
surface of exostome tooth in upper part (2800×); 4 – outer surface of exostome tooth in transition zone (1900×); 5 –
outer surface of exostome tooth in lower part (1900×).
Plagiotheciella Fleisch. ex Broth., nom. il-
leg., was suggested to segregate P. latebricola
and also P. piliferum (and P. passaiensis
(Aust.) Fleisch., which appears to be Isoptery-
giopsis pulchella). However the close relation
of P. piliferum and P. latebricola seems doubt-
ful: among the above listed peculiarities of the
latter species only the low basal membrane is
observed in P. piliferum. Also in this species
the outer surface of exostome is totally papil-
lose (unlike other Altaian species of the genus).
KEY TO THE ALTAIAN SPECIES OF
PLAGIOTHECIUM
1. Plants clearly complanate; leaves asymmet-
ric . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1. Plants not or indistinctly complanate; leaves
±symmetric . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Leaf decurrencies broad, 3-5 cells wide,
composed of ±isodiametric cells; laminal
cells (10-)13-18 μm wide (at 1/3 of leaf
length); capsule curved
. . . . . . . . . . . . . . . . . P. denticulatum (p. 50)
2. Leaf decurrencies narrow, 2-3 cells wide,
composed of elongate cells; laminal cells
5-8 μm wide; capsule nearly straight
. . . . . . . . . . . . . . . . P. laetum (p. 51)
3. Plants small; leaf tips with gemmae (or oth-
erwise at least with some leaves ti ps appear-
ing eroded) . . . . . . . P. latebricola (p. 52)
3. Plants medium-sized; gemmae on abaxial leaf
base, often absent; leaves concave; costa dou-
ble or forked, usually prominent, extending
1/3-4/7 the leaf length
. . . . . . . . . . . . . . . . . . . . . . P. cavifolium (p. 48)
48 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
100 μm
Fig. 23. Plagiothecium denticulatum (Hedw.) B.S.G. (from Kayakkatuyarykskij Creek 1940 m Ignatov 3/152): 1-
4 – leaves; 5 – part of shoot; 6 – habit; 7, 9 – upper lamina cells of stem leaf; 8 – alar cells of stem leaf; 10-11 –
perichaetial leaves; 12 – middle laminall cells of stem leaf; 13 – stem cross-section; 14 – basal leaf cells with rhizoides.
Scale bars: 1 cm – for 6; 2 mm – for 5; 1 mm – for 1-4, 10-11; 100 μm – for 7-9, 12-14.
13 14
15
8
6
34
7
9
10
12
2
1 cm
1 mm
2 mm
11
Plagiothecium cavifolium (Brid.) Iwats.,
J. Hattori Bot. Lab. 33: 360. 1970. – Hypnum
cavifolium Brid., Bryol. Univ. 2: 556 ‘cavifo-
lius’. 1827. Figs. 21 & 22
Plants medium-sized to rather robust, green to
whitish-green, in dense to loose mats. Stem julaceous
to complanate, to 2-3 cm long, with central strand.
Axillar hairs 4-5-celled, 180-230 μm long; apical cell
50-70 × 12-13 μm. Leaves 1.0-2.1 × 0.5-0.8 mm, ovate-
lanceolate, symmetric, with ± equal short decurren-
cies; costa double or sometimes single and forked,
often long, to 1/2 the leaf length; margins recurved
at places. Laminal cells 80-160 × 8-16 (6-14/2) μm;
cells in decurrencies large, thin-walled.
Dioicous. Sporophytes unknown in Altai. SEM
observation of the exostome teeth of a specimen from
Ukrainian Carpatian Mts. (Igantov # MHA) shows
their similarity to P. denticulatum and P. laetum:
teeth ±abruptly narrowed in the transition zone,
subentire above; in the lower part striolate, with
waved cristae; in the middle densely papillose on
few plates; in the upper part above with small low
rare papillae, high and incrassate dorsal trabeculae,
not incrassate margin; ventral surface and trabecu-
lae sparsely spinulose.
Plagiothecium cavifolium is very variable
in Altai, and most of its collections can be sort-
49
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
5
1 2
43
Fig. 24. Plagiothecium denticulatum (Hedw.) B.S.G. (from Kayakkatuyarykskij Creek 1940 m Ignatov 3/152): 1
– peristome (200×); 2 – side view of exostome tooth in upper part, showing papillose inner trabeculae (4000×); 3 –
outer s urface of exostome tooth in upper part (3000×); 4 – outer surface of exostome tooth in transition zone (3500×);
5 – outer surface of exostome tooth in lower part (13000×).
ed into P. cavifolium s. str. (rather small, jula-
ceous plants with very concave leaves, obtuse in
general outline, but often with short reflexed
tips) or P. succulentum (Wils.) Lindb. (medi-
um-sized, ±complanate form with gradually
acuminate leaves). The latter grows in more shad-
ed and wet habitats on soil or rotten wood,
while the former occurs in crevices of ±xeric
rock outcrops. However in deep crevices of rocks
and sometimes in other habitats there are in-
termediates which we were unable to sort to
one of the two. Dimensions of leaves and lami-
nal cells seem correlate with the habit, provid-
ing little help for delimitation of these forms.
Distribution: Widespread species, known from
most European countries, most regions of the former
USSR (from the high Arctic to mountains of Cauca-
sus, Middle Asia, southern Far East) and further to
Himalayas, China, and Japan. In North America it is
equally common in many areas, reaching Georgia,
Alabama, and Arkansas in the south. Also reported
from Antarctic islands and Hawaii. In Altai P. cav-
ifolium grows on rocks and soil at all the altitudes.
Specimens examined: (cavifolium-phenotype): Ayu-
lyuyuzyuk Creek 2050 m (0/393), 2150 (0/130); Bayas
Lake 1750 m (0/1241; 0/657); Bogoyash Creek, in mid-
dle course 2400 m (36/167); Kairu Creek, 16 km up-
stream 1400 m (14/53); Kamga Creek, 14 km upstream
660 m (0/1243); Kobiguayuk Creek 2350 m (0/1240);
Saluru Creek, in middle course 2050 m (36/289); Yailyu
440 m (0/658; 0/659), 460 m (0/1242);
(succilentum-phenotype and intermediate forms): Kish-
te 440 m (Zolotukhin 11.VII.1988); Bolshoye Istyube Creek
470 m (0/1244; 18/18a; 18/56); Chiri Creek, 0.5 km
upstream 450 m (17/110); Karakem River, 3 km upstream
1600 m (0/1415); Chulcha River, in middle course 1100
m (9/75); Elekmonar Creek, 5 km upstream 700 m (26/
50 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
100 μm
Fig. 25. Plagiothecium laetum B.S.G. (from Kamga Creek, 13 km upstream 630 m Ignatov 0/1042): 1 – habit; 2-
5 – leaves; 6 – upper laminal cells; 7 – stem cross-section; 8 – middle laminal cells; 9-10 – perichaetial leaves; 11-
12 – alar cells. Scale bars: 1 cm – for 1; 1 mm – for 9-10; 0.5 mm – for 2-5; 100 μm – for 6-8, 11-12.
1
5
8
6
34
7
910
11
12
2
1 cm 0.5 mm
1 mm
42; 26/52); Kayakkatuyarykskij Creek 1760 m (8/13),
1950 m (8/302); 2050 m Kobukhta 700 m (0/248);
Maloye Istyube Creek, 1 km upstream 700 m (Zolotukhin
22.VII.1991); (7/30); Tura Creek, in middle course 1300
m (28/36); Yurga 450 m (21/12).
Plagiothecium denticulatum (Hedw.) B.S.G.,
Bryol. Eur. 5: 190. 1851. – Hypnum denticulatum
Hedw., Sp. Musc. 237. 1801. Figs. 23 & 24.
Plants medium-sized, in green to dark-green
slightly glossy mats. Stem irregularly branched, to 2
cm long, strongly complanate, with central strand;
epidermal cells thin-walled, not enlarged. Axillary
hair 4-5-celled, 200-270 μm long; apical cell 60-70 ×
12-13 μm. Leaves spreading, 1.5-3 × 0.9-1.1 mm, ovate,
asymmetric, with broad uneven decurrencies, the
upper decurrency wider, 60-65 μm (3-5 cells) wide;
costa double, to 1/3-1/2 the leaf length; margins
plane or recurved, entire or serrulate in the upper 1/
5. Laminal cells 110-140 × 13-18 (12-17/1) μm. In
the decurrencies cells 30-35 × 17-25 μm.
Autoicous. Sporophytes frequent. Perichaetial
leaves 2.1-2.3 × 1 mm, smooth, costa double. Seta
1.5-2.5 cm. Capsule 1.5-2.0 mm long, arcuate, smooth
or slightly furrowed. Operculum obliquely rostrate.
Exostome teeth pale, 0.40-0.48 mm long, abruptly
narrowed in the transition zone, serrulate to nearly
entire in the upper third; in the lower part striolate,
with perfect cristae; in the middle densely papillose
on few plates; in the upper part with low small rare
papillae; dorsal trabeculae prominent, with more high
conic-spinulose papillae; ventral surface and trabe-
culae rugose, with sparse spinulose papillae. Endos-
tome 0.45 mm long, basal membrane 0.14 mm high;
cilia appendiculate. Spores 10-13 μm.
Distribution: The species of nearly cosmopoli-
tan distribution, known in most of the regions of the
Holarctic (being especially common in boreal for-
ests), deeply penetrating to the Arctic, and reoprted
far to the south in Macaronesia, South Africa, Aus-
tralia, and Antarctic islands. In Altai P. denticula-
51
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1
2
3
Fig. 26. Plagiothecium laetum B.S.G. (from Karekem River 1600
m Ignatov 0/1043): 1 – peristome (250×); 2 – side view of exos-
tome tooth in upper part, showing weakly papillose inner trabeculae
(3700×); 3 – outer surface of exostome tooth in transition zone (3300×).
tum is common in mixed and conifer forests grow-
ing on trunk bases, rotten wood, rocks and on soil;
in the alpine zone it grows mostly among big rocks
in rock-fields and as a minor component on soil
banks along stream beds and so on.
Specimens examined: Ayukol 1350 m (0/1249); Ayu-
lyuyuzyuk Creek 2450 m (0/167); Bayas Lake 1750 m
(0/2093a); Bolshoye Istyube Creek 470 m (18/60);
Bolshoye Kurkure Creek, upper course 2300 m (Galanin
9.VIII.1976); Itykul Lake 1800 m (Makhatkov & al.
2.VIII.1978); Kairu Creek, 8 km upstream 1000 m (15/
113); Kairu-Bazhi Peak 2050 m (13/110); Kamga Bay 440
m (Zolotukhin 20.X.1988); Kamga Creek, 13 km upstream
600 m (0/101); Karakol Lakes 1900 m (26/126); Kay-
akkatuyarykskij Creek 1920 m (3/261; 3/270), 1940 m
(3/152); Kishte 440 m (Zolotukhin 11.VII.1988); Kobigu-
ayuk Creek 2200 m (0/1247), 2300 m (0/307; 0/662);
Lesnoi Creek, upper course 2600 m (Zolotukhin 1.VII.1990);
Tamanel Peak 2150 m (34/89); Trekhglavaya Peak 2850
m (Zolotukhin 19.VII.1990); Uedinennoye Lake 880 m (0/
1245); Yailyu 440 m (0/1246), 450 m (0/661).
Plagiothecium laetum B.S.G., Bryol. Eur.
5: 185. t. 495. 1851. Figs. 25 & 26
Plants small to medium-sized, in green to whit-
ish-green glossy mats. Stem irregularly branched, to
1.5 cm long, strongly complanate, with central strand
and non-enlarged, thin-walled epidermal cells. Ax-
illary hair 3-4-celled, 150-200 μm long, apical cell
45-55 × 10-11 μm. Leaves 0.9-1.3 × 0.3-0.4 mm, ovate,
usually recurved nearly throughout, asymmetric, with
unequal decurrencies, the upper decurrency wider,
40-45 μm wide. Laminal cells 80-120 × 5-7 (4-6/1)
μm. In the decurrencies cells rectangular, ca. 50 × 12
μm, in 3-4 rows.
Autoicous. Sporophytes frequent. Perichaetial
leaves 0.9-1.1 × 0.5 mm. Seta 1.5 cm. Capsule 1-1.5
mm long, slightly arcuate to straight. Operculum
obliquely rostrate. Exostome teeth pale, 0.40-0.45
mm long, abruptly narrowed in the transition zone,
serrulate to nearly entire in the upper third; in the
lower part striolate, with perfect cristae; in the
middle densely papillose on few plates; in the up-
per part with low small rare papillae; dorsal trabe-
culae prominent; ventral surface and trabeculae
rugose, with sparse small papillae. Endostome 0.4
mm high, basal membrane ca. 0.07 mm high; seg-
ments narrow and not perforated; cilia nodose.
Spores 8-13 μm.
Though there are many characters to distin-
guish Plagiothecium laetum and P. denticula-
tum, some forms are difficult to interpret, since
they have narrow decurrencies, but wide lami-
nal cells. The latter character seems more reli-
able in such cases.
Distribution: Plagiothecium laetum is a bi po-
lar disjunct species, very common throughout the
boreal zone of the Northern Hemisphere, and spo-
radic also in the Arctic and temperate zones. In Al-
tai it is confined to the forest zone and has never
52 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
100 μm
Fig. 27. Plagiothecium latebricola B.S.G. (from Bolshoye Istyube Creek 470 m Ignatov 18/15): 1 – habit; 2-5 –
leaves; 6 – stem cross-section; 7 – gemmae; 8 – cells of leaf base; 9 – middle laminal cells; 10 – upper laminal cells;
11-12 – perichaetial leaves. Scale bars: 5 mm – for 1; 1 mm – for 11-12; 0.5 mm – for 2-5; 100 μm – for 6-10.
1
5
8
6
3
4
7
9
10
11 12
2
5 mm 0.5 mm
1 mm
been found above the tree-line. It grows on trunks
and rotten wood, but only occasionally can be found
on soil (where P. denticulatum is not rare).
Specimens examined: Ayukol 1000 m (0/1040); Ayu-
lyuyuzyuk Creek 2100 m (0/384); Bayas Lake 1800 m
(0/663); Bolshoye Istyube Creek 470 m (0/1248; 18/
19a); Kairu Creek, 16 km upstream 1400 m (14/42); Ka-
mga Bay 440 m (Zolotukhin 20.X.1988); Kamga Creek,
13 km upstream 600 m (0/1044), 630 m (0/1042); Kar-
akem River, 6 km upstream 1600 m (0/1043); Karakol
Lakes 1900 m (28/149); Kayakkatuyarykskij Creek 1760
m (8/222), 1900 m (8/36), 1920 m (3/160), 1950 m (8/
85), 2000 m (3/86); Kyga River, 2 km upstream 600 m
(0/1041); Oimok 1100 m (Zolotukhin 25.VIII.1986);
Saluru Creek, upper course 2250 m (36/112); Srednij
Shaltan Creek 620 m (0/1045); Tura Creek, in middle
course 1300 m (28/29; 28/49); Yailyu 460 m (0/662),
480 m (0/1046).
Plagiothecium latebricola B.S.G., Bryol.
Eur. 5: 184. t. 494. 1851. – Philoscia latebricola
(B.S.G.) Berk., Handb. Brit. Moss. 146. 1863.
Figs. 27 & 28
Plants small, up 1 cm long, in green glossy
mats. Stem irregularly branched, complanate.
Leaves 0.6-1.0 × 0.2-0.4 mm, ovate-lanceolate,
slightly asymmetric, the upper decurrency (1-)2-
3-cell and 25-40 μm wide. Laminal cells 90-130 ×
5(4/1) μm, in decurrencies ca. 48 × 16 mm. Gem-
mae in leaf axils and on leaf ti ps, 3-4-celled, 51-68
× 11-12 μm.
Dioicous. Sporophytes rather frequent. Pericha-
etial leaves 0.7-0.8 × 0.2-0.4 mm. Seta 1.0 cm. Cap-
sule ±erect, 1.0 mm long. Exostome teeth pale, 0.35
mm long, abruptly narrowed in the transition zone,
serrulate to nearly entire in the upper third; in the
lower part ±striolate, with cristae short, flexuose and
intermixed with papillae; in the middle cristae be-
coming shorter and oblique to longitudinal; in the
upper part with low small papillae arranged in lon-
gitudinal rows; dorsal trabeculae indistinct, with non-
incrassate margin; ventral surface and trabeculae
slightly rugose, without papillae. Endostome 0.3 mm
long, basal membrane low; cilia short or reduced.
Spores ca. 10 μm.
This species is easily recognized by the rel-
atively small size and eroded leaf tips (gem-
mae themselves are not common). Crundwell
(1979) reported cauline rhizoids in P. latebri-
cola, whereas leaf-borne in other Plagiotheci-
um species. According to our observation most
rhizoids in P. latebricola have the same posi-
tion as in other species of the genus and stem-
53
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1 2
43
Fig. 28. Plagiothecium latebricola B.S.G. (from Bolshoye Istyube Creek 470 m Ignatov 18/15): 1 – peristome
(360×); 2 – outer surface of exostome tooth in transition zone (3200×); 3 – upper part of exostome tooth (3000×); 4
– outer surface of exostome tooth in lower part (6000×).
borne only in proximal branches, agreeing with
the pattern observed in other species of Pla-
giothecium.
Distribution: Plagiothecium latebricola is
a rather rare species in most areas where it is
known, at least in the territory of the former
USSR. However it has been found in most of
carefully explored boreal regions. It si also
known from most of the European countries,
Turkey and North America. As some other epix-
ylic boreal species, P. latebricola does not pen-
etrate into the Arctic, or to areas dominated by
steppes, and also is rare in broad-leaved forests.
In Altai P. latebricola was collected on strong-
ly rotten wood in a wet shady canyon rich in
decaying wood.
Specimens examined: Bolshoye Istyube Creek 470 m
(0/1250; 18/15).
Herzogiella Broth.
This genus is characterized by thin-wall epi-
dermal cells and leaves non-decurrent and ser-
rate throughout. However, the type of Herzo-
giella, the South American H. boliviana, may
not be very close to species of the Northern
Hemisphaere (Buck & Ireland, 1985). If further
studies find them different, the correct generic
name for the Holarctic species will be Sharpiel-
la Iwats., J. Hattori Bot. Lab. 28: 202. 1965. The
latter group includes about 5 species, in the
boreal and temperate zones.
In Herzogiella ascendens (Lindb.) Iwats. et
Schof., a North-Pacific species, rhizoids are clus-
tered just below the leaf insertion, pseudopara-
phyllia present and branching is regularly pin-
nate, so it seems closer to Hypnum sect. Breidleria.
54 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 mm
1 cm
100 μm
4
13
1
7
10
23
5
912
811
6
2 mm
Fig. 29. Herzogiella seligeri (Brid.) Iwats. (from Kazakh Altai, Riddera, Polyakov 371): 1-4 − leaves; 5 − capsule; 6
− habit; 7 − shoot; 8 − upper laminal cells; 9 − basal cells; 10 − middle laminal cells; 11-12 − perichaetial leaves; 13 −
stem cross-section. Scale bars: 1 cm − for 6; 2 mm − for 5, 7; 1 mm − for 1-4, 11-12; 100 μm − for 8-10, 13.
Herzogiella seligeri (Brid.) Iwats., J. Hat-
tori Bot. Lab. 33: 374. 1970 – Leskea seligeri
Brid., Musc. Recent. 2(2): 47. 1801 – Sharpiel-
la seligeri (Brid.) Iwatsuki, J. Hattori Bot. Lab.
28: 203. 1965. Figs. 29 & 30
Plants medium-sized, in glossy green or whit-
ish-green tufts. Stem prostrate, rarely irregularly
branching, to 2 cm long, with central strand; epi-
dermal cells thin-walled, small. Axillary hairs 3-4-
celled, 70-90 μm long, apical cell 30 × 8 μm. Rhiz-
oids situated usually in between leaves, not strict-
ly above the center of the leaf below, rarely in more
clear rows above the leaf axil. Leaves spreading, fal-
cate, ca. (1.0-)1.2-2.0 × 0.4-0.7 mm, lanceolate;
costa double; margins serrate above, serrulate be-
low to the base. Laminal cells 70-95 × 7-6 (5-6/1)
μm; at base shorter, in leaf corners practically
not differentiated.
Autoicous. Sporophytes frequent. Perichaetial
leaves 1.3-1.5 × 0.4 mm. Seta to 2.2 cm long. Cap-
sule 2-2.3 mm long, curved and horizontal. Opercu-
lum conic. Annulus of large separating cells. Exos-
tome teeth ca. 0.40 mm long, ±abruptly tapered in
the transition zone, entire; in the lower part imper-
fectly striolate (cristae smooth); in the middle dense-
ly papillose (papillose zone of few plates); in the
upper part with irregularly scattered small conic-
spinulose papillae; dorsal trabeculae and non-in-
crassate margin with conic papillae; ventral sur-
face and trabeculae with rare spinulose papillae.
Endostome 0.40 mm long, basal membrane 0.20 mm
long, segments not perforate; cilia nodose (rarely
appendiculate). Spores 14 μm.
Distribution: In both Eurasia and North Amer-
ica, Herzogiella seligeri occurs mostly in western
regions: in America – from British Columbia to
Oregon, eastward to Montana and Idaho; in Eur-
asia – in most countries (especially abundant in
beech forests), eastward to Urals and Caucasus. In
Asia H. seligeri is known from a few localities in
China (Redfearn & al., 1996) and in Kazakh Altai,
close to the border of Russian Altai, and probably
will be found in the western part of the latter. All
collections were made in Abies forests.
Specimen examined: Kazakh Altai, Riddera, Polyakov 371.
55
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1
3 4
2
Fig. 30. Herzogiella seligeri (Brid.) Iwats. (from Moscow Province, Ignatov 4.VII.1987 (MHA)): 1 – peristome
(250×); 2 – inner side of exostome tooth in upper part, showing papillose inner trabecula (4800×); 3 – outer surface of
exostome tooth in upper part (3600×); 4 – outer surface of exostome tooth in transition zone (3600×).
Hypnaceae Schimp.
A big and heterogeneous family of pleuro-
carps, characterized by (1) straight to falcate-
secund leaves with short double costa; (2) alar
cells often differentiated, but not vesiculate-
inflated; (3) pseudoparaphyllia present; (4)
rhizoids inserted below the leaf insertion; (5)
exostome cross-striolate to papillose, with a zig-
zag centre line and well developed ventral tra-
beculae; (6) endostome with ±high basal mem-
brane and broad, keeled, narrowly perforated
segments, and usually with nodose cilia. Some
to nearly all of these main charachters are
present also in various groups of Sematophyl-
laceae, Entodontaceae, Hylocomiaceae, Plagio-
theciaceae, and Amblystegiaceae, so the limits
of the Hypnaceae with these families remain
disputable. In the course of revisions of these
families many genera were transferred to Hyp-
naceae. However these attempts to increase the
naturalness of Sematophyllaceae, Plagiotheci-
aceae, Entodontaceae, etc. have resulted in an
enormous inflation of the Hypnaceae.
The following genera, found in Altai, pose
problems of familial alliance:
Hypnaceae/Hylocomiaceae: Gollania, Rhy-
tidiadelphus, Pleurozium;
Hypnaceae/Plagiotheciaceae: Taxiphyllum,
Isopterygiopsis, Herzogiella, Orthothecium;
Hypnaceae/Entodontaceae: Platygyrium,
[Orthothecium];
Hypnaceae/Sematophyllaceae: Heterophyl-
lium, Struckia;
Hypnaceae/Amblystegiaceae: Campylium
56 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
s. l. (incl. Campyliadelphus and Campylophyl-
lum), Calliergonella, Psuedohygrohypnum.
These genera are discussed in corresponding
genera or families, except the Amblystegiaceae
which will be treated in a separate paper.
Altaian members of the Hypnaceae belong
mostly to the subfamily Hypnoideae as it was
defined by Nishimura & al. (1984). The rela-
tively homogeneous group around Hypnum in-
cludes Callicladium, Eurohypnum, Gollania,
Ptilum, Homomallium, and Heterophyllium. Es-
pecially distinct is Pylaisiella, comprising the
small subfamily Pylaisielloideae. The recently
described Podperaea is referred here to Cten-
idioideae, to which belongs its close relative Mit-
tenothamnium. We place Taxiphyllum in Hyp-
naceae, not in Plagiotheciaceae, however in the
former its position is also rather isolated. Pre-
liminary placement of Taxiphyllum in Hyp-
noideae is probably not exact, and maybe Tax-
iphyllum and Isopterygium s. str. need their
own subfamily.
Subfamily Pylaisielloideae Fleisch.
The subfamily is characterized by straight
leaves, erect capsules with advanced peristome
characters.
Pylaisiella Kindb.
This genus includes ca. 30 species, distribut-
ed in temperate regions, with 2-3 species in the
boreal zone. Pylaisiella differs from other mem-
bers of Hypnaceae in the perfectly straight cap-
sule and in structure of both endostome and
exostome. Cilia of endostome are short and with-
out appendicules. Exostome teeth are (1) heavi-
ly papillose on the outer surface above; (2) dor-
sal trabeculae in proximal tooth high and plates
between them are smooth; (3) the inner layer
of the teeth (derived from PPL) is thick due to
deposition of additional material. These pecu-
liarities cause anomalous hygrocastique hygro-
scopic movements – the teeth are ±involute
when dry and become erect to somewhat re-
flexed when wet (this character seen in P.
polyantha, P. selwynii, P. brotheri and P. in-
tricata). In this respect Pylaisiella differs from
most boreal pleurocarps, and resembles many
tropical mosses and, in the northern bryoflora,
Neckera, Leucodon, Homalothecium, Entodon,
Anomodon, and Leskeaceae (Leskea, Iwatsuk-
iella, Myrinia, etc.). These genera disperse spores
in wet periods (usually in the cool half of the
year) and keep capsules ±closed in dry peri-
ods. This pattern can be assumed to be an adap-
tation to the true epi phytic life (Mueller &
Neumann, 1988). However, the pattern of hy-
groscopic movements is hardly enough to seg-
regate Pylaisiella in its own family, since this
character is variable within many families: Neck-
eraceae (xerocastique, or involute when wet in
Homalia/ hygrocastique in Neckera & many
other genera), Brachytheciaceae (hygrocastique
in Homalothecium and some Rhynchostegiel-
la/ xerocastique in many other genera), etc.
KEY TO THE ALTAIAN SPECIES OF
PYLAISIELLA
1. Branches strongly curved; alar cells numer-
ous, to 25 along the margin; capsule ovoid;
exostome and endostome tightly attached
to each other, and fused below; spores 18-
30 μm . . . . . . . . . . . . . . . . . . . . . . . . P. selwynii
1. Branches straight to little-curved; alar cells
less numerous, to 15 along the margin; cap-
sule ±cylindric; exostome and endostome not
attached to each other; spores 12-19 μm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. polyantha
Pylaisiella polyantha (Hedw.) Grout, Bull.
Torrey Bot. Club 23: 229. 1896. – Leskea poly-
antha Hedw., Sp. Musc. 229. 1801. Figs. 31 & 32
Plants medium-sized, yellow-green to golden-
brownish, glossy. Stem creeping, 4 cm long, ±regular-
ly branched, branches 5-6 mm long; tips of stem and
branches often ±arcuate; epidermal cells of stem
thick-walled; central strand well-developed.
Pseudoparaphyllia narrowly triangular, strongly ser-
rate to incised. Axillary hairs 4-celled, 100-110 μm
long, apical cell 40-50 × 9-10 μm. Stem leaves straight,
1.2-1.5 × 0.4-0.5 mm, ovate-lanceolate; margins plane,
entire. Branch leaves 1.0-1.1 × 0.3-0.4 mm. Laminal
cells 35-70 × 5-7 (4-5.5/1-1.5) μm. Alar cells sub-
quadrate, forming a triangular indistinctly delimit-
ed group, ca. 15 along the margin.
Autoicous. Sporophytes frequent. Perichaetial
leaves 1.4-1.5 × 0.4-0.5 mm, smooth. Seta ca. 1 cm.
Capsule 1.5 mm long, straight, cylindric. Operculum
conic. Annulus of 3 rows of rectangular cells, partly
deciduous, but almost indistinctly delimited from
the lower cells. Exostome teeth 0.3 mm long, gradu-
ally tapered, entire, on the outer surface with high
trabeculae. Dorsal trabeculae smooth in proximal
tooth, in the upper 2/3 papillose and incrassate;
papillae large, high and often forked. Plates in be-
tween trabeculae moderately roughened with low
round papillae to rugose. Ventral trabeculae heavily
57
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
100 μm
Fig. 31. Pylaisiella polyantha (Hedw.) Grout (from Adylda Creek 1100 m Ignatov 34/130): 1 – upper laminal
cells of branch leaf; 2 – upper laminal cells of stem leaf; 3-5 – stem leaves; 6 – habit; 7-8 – branch leaves; 9-11 –
perichaetial leaves; 12 – capsule; 13 – middle laminal cells of stem leaf; 14 – pseudoparaphyllium; 15 – stem cross-
section; 16 – alar cells of stem leaf; 17 – alar cells of branch leaf. Scale bars: 5 mm – for 6; 2 mm – for 12; 1 mm –
for 9-11; 0.5 mm – for 3-5, 7-8; 100 μm – for 1-2, 13-17.
17
15
16
13 14
5 mm
1
5
8
6
3
4
7
910 11
12
0.5 mm
2
1 mm 2 mm
incrassate, rugose. Endostome free, 0.35 mm long, with
basal membrane ca 0.12 mm high, segments ±papil-
lose above; cilia short to absent. Spores 12-19 μm.
Distribution: This species is widely distributed
throughout the Holarctic, from the northern limit of
trees to the subtropical zone. In Altai P. polyantha
never exceeds tree-line. Collection were made from
trunks and twigs (Populus, Salix, Betula, Sorbus,
Padus, Abies) at lower elevations, and sometimes
from rotten logs and stumps. Above 1400 m it grows
mostly on rocks, and bases of shrubs of Caragana
and Rhododendron.
58 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 2
43
Fig. 32. Pylaisiella polyantha (Hedw.) Grout (from Bele 500 m Ignatov 0/79): 1 – peristome (330×); 2 – outside
view of exostome teeth (1150×); 3 – outer surface of endostome in the middle (4250×); 4 – side view of upper part of
exostome tooth, showing strongly incrassate inner trabeculae (5700×).
Specimens examined: Adylda Creek 1100 m (34/130; 34/
82); Akkurum 550 m (Zolotukhin 22.IX.1989); Ayukol 950 m
(0/1199); Bele 500 m (0/79); Berekhtuyaryk 1600 m (0/
1695), 1640 m (0/608); Bolshoi Yaloman Creek, in middle
course 1500 m (Makarov & al. 15.VII.1972); Chemal, 5 km to
the SE 500 m (Makarov 28.VI.1972); Chiri Creek, 0.5 km
upstream 450 m (17/22; 17/87); Chodro 900 m (Lebedeva &
al. 25.VIII.1979); Chulcha River, in middle course 1000 m (9/
47); Elekmonar Creek, 5 km upstream 700 m (26/38; 26/43);
Kamga Creek, 2 km upstream 450 m (Zolotukhin 20.X.1988);
Kamga Creek, at mouth 440 m (Zolotukhin 20.X.1988); Kar-
akem River, 6 km upstream 1600 m (0/1195); Karatyt 1300 m
(Zolotukhin 20.II.1990); Kayakkatuyarykskij Creek 1850 m
(8/293), 1940 m (3/246); Korbu 440 m (0/1197), 450 m
(Zolotukhin 20.X.1988); Kukol 1800 m (0/609); Maima 280
m (35/23); Malyi Yaloman 1100 m (25/6; 25/88); Malyi
Yaloman Creek, 8 km upstream 1100 m (25/19); Srednij Shal-
tan Creek 640 m (0/1196); Ust-Sema 400 m (24/160); Yai-
lyu 440 m (0/610), 450 m (Zolotukhina 14.VII.1988).
Pylaisiella selwynii (Kindb.) Crum, Steere
et Anderson, Bryologist 67(2): 164. 1964. – Py-
laisia selwynii Kindb., Ottawa Naturalist 2:
156. 1889. – Pylaisia schimperi Card., Bull. Herb.
Boiss. 7: 373. 1899. Figs. 33 & 34
Plants medium-sized, yellow-green, glossy. Stem
creeping, 5 cm long, ±regularly branched, branches
5-6 mm long, tips of stem and branches often arcu-
ate to circinate when dry; epidermal cells of stem
thick-walled; central strand ill-developed.
Pseudoparaphyllia narrowly triangular, strongly ser-
rate to incised. Axillary hairs 3-celled, 100-110 μm
long, apical cell 65 × 10 μm. Stem leaves straight, 1.0-
1.2 × 0.4-0.5 mm, ovate-lanceolate; margins plane,
entire. Branch leaves 0.8-0.9 × 0.2-0.3 mm. Laminal
cells 30-55 × 7-8 (3.5-4.5/3-3.5) μm. Alar cells sub-
59
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 33. Pylaisiella selwynii (Kindb.) Crum & al. (from Kamga Creek, 2 km upstream 450 m Zolotukhin 20.X.1988):
1 – capsule; 2 – habit; 3 – stem cross-section; 4-5 – stem leaves; 6-7 – branch leaves; 8 – upper laminal cells of
branch leaf; 9 – upper laminal cells of stem leaf; 10 – alar cells of stem leaf; 11 – alar cells of branch leaf; 12 –
middle laminal cells of stem leaf; 13 – middle laminal cells of branch leaf; 14-15 – pseudoparaphyllia; 16-18 –
perichaetial leaves. Scale bars: 5 mm – for 1-2; 1 mm – for 4-7; 0.5 mm – for 16-18; 100 μm – for 3, 8-15.
17
15 16
14
1
58
6
3
4
7
9
10
12
2
18
5 mm
0.5 mm
100 μm
1 mm
11 13
quadrate, forming large indistinctly delimited group,
reaching ca. 25 along the margin.
Autoicous. Sporophytes frequent. Perichaetial
leaves 1.9-2.0 × 0.5-0.6 mm, entire; costa double, short
to long, reaching mid-leaf. Seta ca. 1 cm. Capsule 1.5
mm long, ovoid, straight, pale and often contrasting-
ly colored below the mouth. Operculum conic. An-
nulus of 2-4 rows of quadrate and transversely rect-
angular cells, partly deciduous, but indistinctly de-
limited from the lower cells. Exostome teeth 0.25
mm long, gradually tapered, entire, with high dorsal
trabeculae. Dorsal trabeculae smooth below, papil-
lose in the upper 1/2-1/3. Plates smooth below, with
round low papillae above (of the same type as on the
dorsal trabeculae). Ventral surface and high ventral
trabeculae with somewhat larger dense papillae. En-
dostome as high as the teeth, in the lower 1/3 adher-
ent to the latter, in the upper part densely papillose,
perforated and easily splitting off along the keel.
Spores 20-32 μm, roughly papillose.
More strongly curved shoots and ovoid (not
cylindric) capsules allow differentiation of this
species from Pylaisiella polyantha in the field
with certainty.
Distribution: The distribution of Pylaisiella sel-
wynii was mapped and discussed by Ignatov (1992).
The species is common in East Asia and eastern North
America, with limited occurrence in western sectors
of both continents. In Altai (as well as in all other
parts of Russia) this species grows exclusively as an
epiphyte, mostly on Populus tremula, rarely on P.
laurifolia, Salix, Betula, and confined to only lower
elevations below 600 m. On Populus tremula it grows
to 10 m and more above the ground, especially if the
trunk is somewhat inclined. Often in such cases P.
selwynii is the only epiphyte species and forms pure
60 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1
2
4
5
3
Fig. 34. Pylaisiella selwynii (Kindb.) Crum & al. (1, 4
– from Kamga Creek 460 m Ignatov 0/1202; 2, 3, 5 –
from Chulyshman River 500 m Zolotukhin 23.IX.1988):
1 – peristome (430×); 2 – outer surface of exostome teeth
(900×); 3 – side view of exostome tooth in upper part,
showing coarsely papillose both surfaces and trabeculae
(4650×); 4 – part of peristome, showing densely papillose
exostome segments (1200×); 5 – outer trabeculae at about
two-thirds of exostome tooth (5500×).
strips on the overhanging surface of the trunk (the
mainstream of rain-water). In Gorno-Altaiask P. sel-
wynii tolerates strong air pollution, occurring in the
central part of the city on trunks of Populus with P.
polyantha, Ceratodon purpureus (Hedw.) Brid., etc.
Specimens examined: Akkurum 500 m (Zolotukhin
21.IX.1989); Balykcha 450 m (Schischkin 6.IX.1931 LE!);
Chemal Creek, 3 km upstream 450 m (34/135); Chulysh-
man River, at Bashkaus mouth 500 m (Zolotukhin
23.IX.1989); Gorno-Altaisk 400 m (35/16); Gorno-Al-
taisk (SE edge) 600 m (0/1198; 23/5); Kamga Creek, 2
km upstream 450 m (Zolotukhin 20.X.1988; 0/1201);
Kamga Creek, 3 km upstream 460 m (0/1202); Korbu
440 m (Zolotukhin 20.X.1988); Kyrsai 450 m (Schisch-
kin 4.IX.1931 LE!); Pyankovo (Vereshchagin 11.VI.1905
LE!); Yailyu 440 m (Zolotukhin 6.VIII.1988).
Subfamily Ctenidioideae Fleisch.
The subfamily is characterized by tenden-
cies to the marked differentiation of stem and
branch leaves, non-falcate leaves, distinct leaf
serration, prorate laminal cells, and weakly dif-
ferentiated alar cells.
Podperaea Iwats. et Glime
This monotypic genus is distinct in having
(1) strongly enlarged epidermal cells; (2) dis-
tinctly prorate laminal cells; (3) “double-celled”
teeth at leaf margins formed by the curved
upper end of lower cell and the curved lower
end of upper cell. Iwatsuki & Glime (1984)
61
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 35. Podperaea krylovii (Podp.) Iwats. et Glime (from Kobukhta 460 m Ignatov 0/626): 1-3 – stem leaves; 4-
5 – branch leaves; 6-7 – perichaetial leaves; 8 – upper laminal cells of stem leaf; 9 – alar cells of stem leaf; 10 –
middle laminal cells of stem leaf; 11 – stem cross-section; 12 – pseudoparaphyllia; 13-14 – habit; 15 -capsule. Scale
bars: 1 cm – for 13; 2 mm – for 14-15; 0.5 mm - for 1-5; 1 mm – for 6-7; 100 μm – for 8-12.
15
13
14
15
8
6
3
4
7
9
10
11
12
100 μm
2
1 cm
0.5 mm
1 mm
2 mm
compared this genus with Mittenothamnium,
which is close to Podperaea in prorate cells
and some other details (general appearance,
capsule shape, peristome characters), but differs
in characters 1 & 3.
Podperaea krylovii (Podp.) Iwats. et Glime,
J. Hattori Bot. Lab. 55: 495. 1984. – Chrysohyp-
num krylovii Podp., Spisy Prir. Fak Masaryk-
ovy Univ. 116, t. 28, f. 19 ‘krylovi’. 1929. – Campy-
lium krylovii (Podp.) Lazarenko, J. Bot. Acad.
Sci. Ukraine 2: 288. 1941. Figs. 35 & 36
Plants small, green, forming loose mats or growing
as solitary shoots among other mosses. Stem pros-
trate, ca. 1 cm long, with markedly differentiated hy-
alodermis, without central strand, irregularly branched,
branches 2-4 mm long. Pseudoparaphyllia narrow-lan-
ceolate. Axillar hairs 3-celled, 65 μm long, apical cell
25 × 8 μm. Stem leaves widely spreading, 0.4-0.8 × 0.2-
0.3 mm, ovate-lanceolate, gradually acuminate, serrate
all around, the teeth near the base “double-celled”;
costa double or forked, reaching 1/4-1/2 the leaf
length. Branch leaves 0.4-0.5 × 0.1-0.2 mm. Laminal
cells 30-55(-75) × 5 (4/1) μm, with papillae on the
upper and rarely also the lower ends (Fig. 36-7). Alar
cells few, enlarged, indistinctly delimited.
Autoicous. Sporophytes frequent. Perichaetial
leaves 0.9 × 0.2 mm, ecostate, serrulate, smooth. Seta
1.7 cm. Capsule ca. 1 mm long, arcuate and strongly
contacted below the mouth when dry. Operculum
conic. Annulus not differentiated. Exostome teeth ca.
0.4 mm long, abruptly tapered in the transition zone,
serrate in the upper third; striolate below; in the
middle few plates densely papillose; in the upper
part sparsely papillose with small low irregularly
arranged papillae; dorsal trabeculae low; ventral
trabeculae smooth or with rare spinulose papillae.
Endostome 0.5 mm long, basal membrane 0.2 mm
high; cilia 2-3, nodose. Spores 10-12 μm.
Podperaea is easy to recognize by “double-
celled” teeth of leaf margins. Superficially it is
similar to Campylium sommerfeltii Myr.
Distribution: The distribution of this species was
mapped by Ignatov (1990), but some new informa-
tion has appeared since that time. The record from
Chukotka Peninsula was found to be erroneous (the
collection in LE is a mixture of Campylium som-
merfeltii and Myurella tenerrima), and also Pod-
peraea have been found in Liaoning Province of
China (Buck 23759A, NY!). So, now its range includes
South Siberia (westward to Altai), Russian Far East,
northern Japan (Hokkaido) and northeastern China.
62 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1
7
2
4 5
6
3
Fig. 36. Podperaea krylovii (Podp.) Iwats. et Glime (from Kobukhta 460 m Ignatov 0/626): 1 – peristome (230×);
2 – front view of upper part of exostome tooth (2600×); 3 – side view of upper part of exostome tooth (2300×); 4 –
outer surface of exostome tooth in transition zone (3400×); 5 – outer surface of exostome tooth in lower part (11000×);
6 – outer surface of exostome tooth below transition zone (4300×); 7 – surface of stem leaf (2850×).
In Altai Podperaea has been found around Teletz-
koye Lake, but locally is rather common here. It grows
on wet rich soil in ravines, open eroded slopes and
occasionally also on rather dry rotten logs.
Specimens examined: Bele 530 m (0/1192); Kobukhta 460 m
(0/626); Yailyu 450 m (1/10; 1/40; 1/68), 480 m (0/1191).
63
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 37. Homomallium connexum (Card.) Broth. (1-15 – a slender form, from Baigazan 450 m Ignatov 19/3; 16-21
– a robust form, from Kyrsai 450 m Bardunov 6.VII.1966): 1-3, 16-18 – leaves; 4-7 – perichaetial leaves; 8, 20 –
middle laminal cells; 9-10 – pseudoparaphyllia; 11, 21 – alar cells; 12, 19 – upper laminal cells; 13 – stem cross-
section; 14 – capsule; 15 – habit. Scale bars: 5 mm – for 15; 2 mm – for 14; 1 mm – for 4-7; 0.5 mm - for 1-3, 16-
18; 100 μm – for 8-13, 19-21.
5 mm
2 mm
100 μm
1 mm
0.5 mm
12
5
8
6
23
47
9
1
10
11 13
15
14
17
21
20
18 19
16
Subfamily Hypnoideae
The subfamily is characterized by a tenden-
cy toward falcate-secund leaves, smooth cells,
differentiated alar cells, inclined and ±curved
capsule, and normal hypnoid peristome. How-
ever, since the subfamily is large there are excep-
tions: leaves are straight in Callicladium, Eu-
rohypnum, Homomallium, Heterophyllium, and
Taxiphyllum; cells are prorate in some Golla-
nia; alar cells are weakly differentiated in Tax-
iphyllum and some species of Hypnum.
Homomallium (Schimp.) Loeske
The genus has about 10 species, distributed
in temperate regions. Brotherus (1925) placed
64 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 38. Homomallium incurvatum (Brid.) Loeske (from Kobukhtushka 440 m Ignatov 0/634): 1 – habit; 2 –
capsule; 3-4 – leaves; 5-6 – perichaetial leaves; 7, 10 – upper lamina cells; 8 – stem cross-section; 9 – pseudopara-
phyllium; 11-12 – middle lamina cells; 13-14 – alar cells. Scale bars: 5 mm – for 1; 2 mm – for 2; 1 mm – for 5-6;
0.5 mm – for 3-4; 100 μm – for 7-14.
4
3
5
2
6
7
10
9
812
13 14
100 μm11
5 mm
2 mm
1 mm
0.5 mm
Homomallium into the Pylaisioideae due to straight
leaves, without differentiation of stem and branch
leaves. However, inclination of capsule and im-
portant characters of the peristome of Homomal-
lium are just the same as in Hypnoideae, and Ando
(1964) transferred Homomallium in this subfam-
ily. Differences from Hypnum are not many, but
about 10 species of Homomallium form a ±well-
delimited group of small autoicous plants with
straight leaves whose alar cells are homogeneous-
ly subquadrate, forming a distinct area, and cap-
sules relatively long and often distinctly curved
when dry. East Asian species of Homomallium
were revised by Ando (1964, 1965).
KEY TO THE ALTAIAN SPECIES OF HOMOMALLIUM:
1. Leaves broadly ovate-lanceolate, ±short-
acuminate; median laminal cells 20-40 μm
long; alar cells numerous, 20-30 along the
margin . . . . . . . . . . . . . . . . . . . . . H. connexum
1. Leaves narrowly ovate-lanceolate, ±long-
acuminate; median laminal cells 30-60 μm
long; alar cells less numerous to few, 10-20
along the margin . . . . . . . . . H. incurvatum
Homomallium connexum (Card.) Broth.,
Nat. Pfl. 1: 1027. 1908. – Amblystegium con-
nexum Card., Beih. Bot. Centralbl. 17: 39. f.25.
1904. Figs. 37 & 39 (1,3,5)
Plants small, in green to dark-green mats. Stem ca.
1.5 cm long, without central strand, irregularly
branched; branches to 5 mm long. Pseudoparaphyllia
lanceolate. Axillary hairs 2-celled, 40 μm long, apical
cell 25 × 9 μm. Leaves straight, imbricate, ovate-lan-
ceolate, 0.5-1.0 × 0.2-0.4 mm; margins subentire; costa
sometimes lacking to double or forked, 1/3 of leaf length.
Laminal cells 20-40 × 5-6 (4-5/1) μm. Alar cells sub-
quadrate to transverse, 20-30 along the margin.
Autoicous. Sporophytes frequent. Perichaetial
leaves 1.5-1.8 × 0.5-0.6 mm, costa to 1/3-1/2. Seta 1
cm. Capsule ca. 1.5-1.8 mm long, strongly arcuate;
65
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
4
5 6
1 2
3
Fig. 39. Homomallium connexum (Card.) Broth. (1, 3, 5, from Baigazan 450 m Ignatov 19/3) and H. incurvatum (Brid.)
Loeske (2, 4, 6, from Ust-Sema 500 m Ignatov 24/99): 1-2 – peristomes (1 – 230×; 2 – 280×); 3-4 – side views of upper parts
of exostome teeth (3 – 4250×; 4 – 3700×); 5-6 – outer surface of exostome teeth in transition zone (5 – 3500×; 6 – 2600×).
operculum conic-apiculate. Exostome teeth 0.45 mm
long, abruptly tapered in the transition zone, serrate
in the distal third; in the lower part striolate, with
perfect cristae; in the middle few plates densely pap-
illose; in the upper part sparsely small-papillose;
dorsal trabeculae, incrassate margin, ventral surface
and ventral trabeculae with small low papillae. En-
dostome as long as teeth, basal membrane, 0.15 mm
high; cilia appendiculate. Spores 12 μm.
Distribution: This species is known from Japan,
Korea, China (most of the provinces), and Russian Far
East. In Altai it grows in areas with rather mild cli-
mate, on ±exposed rocks. This is the first record in
South Siberia.
66 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 40. Eurohypnum leptothallum (C. Müll.) Ando (1, 7-8 – from Chemal 450 m Ignatov 34/146; 2-6, 9-11 – from
Ust-Sema 340 m Ignatov 24/141): 1, 7 – shoots with perichaetia; 2 – habit; 3-4 – leaves; 5 – stem cross-section; 6
– upper laminal cells; 8 – capsule; 9 – pseudoparaphyllia; 10 – middle laminal cells; 11 – alar cells. Scale bars: 1cm
– for 2; 2 mm – for 8; 1 mm – for 1, 7; 0.5 mm – for 3, 4; 100 μm – for 5-6, 9-11.
2
5
8
9
11
7
6
3
1
4
10
1 cm
1 mm
100 μm
1 mm
0.5 mm
Specimens examined: Baigazan 450 m (19/3); Edikh-
ta Creek 1100 m (34/164); Kyrsai 450 m (Bardunov
6.VII.1966).
Homomallium incurvatum (Brid.) Loeske,
Hedwigia 46: 314. 1907. – Hypnum incurva-
tum Brid., Muscol. Recent. 2(2): 119. 1801.
Figs.38 & 39 (2,4,6)
Plants small, in dark-green mats. Stem ca. 1.5 cm
long, without central strand, irregularly branched;
branches to 4 mm long. Pseudoparaphyllia lanceolate.
Axillary hair 2-celled, 70 μm long, apical cell 60 × 8
μm. Leaves straight, erect, narrowly ovate-lanceolate,
0.6-1.0 × 0.2-0.4 mm, lanceolate, margins entire. Lam-
inal cells 30-60 × 4-5 (3-4/1) μm. Alar cells 10-20
(-25) along the margin.
Autoicous. Sporophytes frequent. Perichaetial
leaves 1.7-1.8 × 0.4-0.5 mm. Seta 1.2 cm. Capsule ca.
2 mm long, strongly arcuate; operculum conic-api-
culate. Exostome teeth 0.55 mm long, abruptly ta-
pered in the transition zone, serrate in the distal third;
in the lower part striolate; in the middle few plates
densely papillose; in the upper part sparsely rough-
ened with small low and often imperfect papillae;
dorsal trabeculae, incrassate margin, ventral surface
and ventral trabeculae with small low papillae. En-
dostome as long as the teeth, basal membrane 0.23
mm high; cilia appendiculate. Spores 13 μm.
Homomallium plagiangium C. Müll., de-
scribed from eastern China, differs from typi-
cal H. incurvatum in more numerous and more
transparent alar cells, and many Altaian speci-
mens fit well in this diagnosis. However, inter-
mediate specimens occur and further study is
necessary to understand if this species is really
distinct from H. incurvatum.
Distribution: Homomallium incurvatum is
known from Europe, Ural Mts., Caucasus, Middle
67
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1 2
3 4
Fig. 41. Eurohypnum leptothallum (C. Müll.) Ando (from Chemal 450 m Ignatov 34/146): 1 – peristome (220×);
2 – outer surface of exostome tooth in upper part (4400×); 3 – side view of exostome tooth in upper part, showing
smooth inner trabeclae (4400×); 4 – outer surface of exostome tooth in transition zone (3400×).
East, Mongolia, China, Japan, and Himalayas. In Al-
tai this species occurs at lower elevations, on dif-
fusely shaded rocks, mostly calcareous.
Specimens examined: Belokurikha 400 m (Bardunov
5.VI.1966); Bijka Creek, 4 km upstream along Katun
450 m (34/166); Chiri 450 m (17/45); Chiri Creek,
0.5 km upstream 450 m (17/88); Chulcha River, in
middle course 1100 m (9/44; 9/67); Kobukhta 550 m
(0/2055); Kobukhtushka 440 m (0/634); Kumalyr
Verkhnij 1040 m (Krylov 22.VI.1903 LE!); Kumzir 450
m (20/24; 20/25); Kyrsai 450 m (a; Bardunov
6.VII.1966); Ust-Sema 400 m (24/97), 450 m (0/1714),
500 m (24/127; 24/99).
Eurohypnum Ando
This monotypic genus resembles Homo-
mallium in straight leaves and indistinctly
plicate perichaetial leaves, but differs in more
robust plants (about the size of Hypnum
vaucheri), often irregularly branched, terete
foliage; leaves distinctly serrulate above, with
extensive alar regions; perichaetial leaves
coarsely serrate in acumen; nearly straight
to slightly inclined capsule (markedly
curved in most of Homomallium, though in
H. mexicanum Card. described as variable
to suberect, cf. Crum & Anderson, 1981); and
dioicous sexual condition. It is also close to
Hypnum, but differs in well-developed, some-
times thick-walled central strand and
straight leaves.
Some older publications refer E. leptothal-
lum to Erythrodontium of Entodontaceae, also
a ±julaceous plant, which however has defi-
nitely straight capsules and reduced peristome.
68 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Eurohypnum leptothallum (C. Müll.) Ando,
Bot. Mag. Tokyo 79: 761. 1966. – Cupressina
leptothalla C. Müll., Nuov. Giorn. Bot. Ital., n. s.,
3: 119. 1896. – Hypnum leptothallum (C. Müll.)
Par., Ind. Bryol. Suppl.: 204. 1900. – E. leptothal-
lum f. tereticaule Nog., J. Jap. Bot. 15: 760. 1939.
Figs. 40 & 41
Plants medium-sized, green to brown-green. Stem
prostrate, 3 cm long, without hyalodermous, with cen-
tral strand; branching strongly depends on light
conditions: in shade, shoots slender, prostrate and
very rarely irregularly branched, while in sunny places,
regularly (though not densely) pinnate-branched,
more stiff, sometimes tumid. Pseudoparaphyllia nar-
row lanceolate. Axillary hairs 8-celled, 170-200 μm
long, apical cell ca. 40 × 10 μm. Stem leaves straight
to somewhat falcate above, 1.0-1.2(-1.5) × 0.4-0.5
(-0.7) mm, ovate, acuminate; margins serrate above,
entire below. Laminal cells 35-55 × 5-6(3-4/2) μm.
Alar cells subquadrate to transversely ovate, form-
ing an indistinctly delimited large group, ca. 25-30
cells along the margin.
Dioicous. Sporophytes found in Altai in only one
collection (calcareous rocks on steep slope to the
Katun River, in diffuse shade of scattered pines; for-
est-steppe zone). Perichaetial leaves 3 mm long, indis-
tinctly plicate, strongly serrate above; costa short and
double, to 1/3 the leaf length. Seta 1.5 cm. Capsule ca.
1.5 mm long. Exostome teeth 0.5 mm long, gradually
narrowed, slightly serrulate above; striolate (cristae
smooth) below; in the middle cristae changing orien-
tation to oblique and longitudinal; in the upper part
longitudinally striolate with cristae low-papillose;
dorsal trabeculae low, with margin slightly incras-
sate; ventral surface and trabeculae smooth. Endos-
tome 0.55 mm long, basal membrane 0.2 mm high;
cilia nodose or reduced. Spores 15 μm.
Eurohypnum is similar to Hypnum vaucheri
in numerous alar cells and concave leaves, but
differs from it by leaves straight and clearly
serrulate to serrate above.
Distribution: The species is common in many
regions of Japan, Korea, China, Taiwan, Mongolia,
Russian Far East and South Siberia. In Altai Euro-
hypnum is common in ±dry habitats within the for-
est and forest-steppe zones, growing mostly on rocks,
very rarely on Populus trunks.
Specimens examined: Aedigan Creek, 5 km upstream
600 m (34/129; 34/43); Artyshtu Creek 600 m (Zolo-
tukhin 20.IX.1989), 650 m (Zolotukhin 20.IX.1989);
Arzhan (Chulyshman lower course) 530 m (Zolotukhin &
al. 24.IV.1978); Chemal 450 m (29/46); Chemal, 10 km
upstream along Katun 450 m (34/146; 34/30); Chiri
450 m (0/1194; 0/2020; 17/63; 17/92); Chiri Creek,
0.5 km upstream 450 m (17/2; 17/6); Chulcha River, in
middle course 820 m (0/633); Chulyshman River, 5 km
upstream 750 m (Zolotukhin & al. 26.IV.1978); Derbogach
450 m (0/24; 0/4); Kuba Creek, 5 km apart from Che-
mal 500 m (Makarov 3.VI.1972); Maima 280 m (35/50;
35/51; 35/52), 350 m (35/9); Malyi Yaloman 900 m
(25/143), 950 m (25/73); Malyi Yaloman Creek, 4 km
upstream 900 m (25/67); Ust-Sema 340 m (24/141), 350
m (24/139), 450 m (24/50), 500 m (0/1193); Yailyu
450 m (1/30; 1/4).
Hypnum Hedw.
A large genus with about 60 species, distrib-
uted in all zones of both hemispheres. The main
diagnostic characters include regularly pinnate
plants, falcate-secund leaves with short double
costa, curved to arcuate capsule, and perfect
peristome. Subdivision of the genus into 11 sec-
tions was suggested by Ando (1973), of which
the monotypic section Polyptera was later seg-
regated as a new genus Caribaeohypnum (Ando
& Higuchi, 1984). Two other sections are some-
times considered as separate genera, Pseudoste-
reodon and Breidleria.
KEY TO THE ALTAIAN SPECIES OF HYPNUM
1. Epidermal cells of stem large, ±thin-walled
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1. Epidermal cells of stem not differentiated,
thick-walled . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2. Plants large (stem leaves 0.7-0.9 mm
wide) . . . . . . . . . . . . . H. pratense (p. 85)
2. Plants small to medium-sized (stem leaves
0.3-0.6(0.7) mm wide) . . . . . . . . . . . . . . 3
3. Plants medium-sized, autoicous, usually with
sporophytes; epidermal cells weakly differ-
entiated . . . . . . . . . . . . . . . H. fauriei (p. 80)
3. Plants small-sized; dioicous, only rarely with
sporophytes; epidermal cells clearly differ-
entiated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Stem leaves triangular-lanceolate, clearly
rounded towards the cordate base; shoots
julaceous to complanate; margins plane;
plants densely plumose-branched
. . . . . . . . . . . . . . . . . . H. plicatulum (p. 85)
4. Stem leaves ovate-lanceolate, weakly nar-
rowed to the insertion; margins sometimes
recurved below and incurved in transi-
tion to acumen, so the upper part is tubu-
lose; shoots clearly julaceous; plants reg-
ularly to ±irregularly branched
. . . . . . . . . . . . . . . . . . H. hamulosum (p. 83)
5. Stems regularly pinnate-branched; plants au-
toicous, usually bearing sporophytes . . . 6
69
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 42. Hypnum recurvatum (Lindb. & H. Arnell) Kindb. (from Bayas Lake 1860 m Ignatov 0/1722): 1-3 – stem
leaves; 4 – middle laminal cells of stem leaf; 5-7 – branch leaves; 8 – habit; 9-10 – alar cells of stem leaf; 11 – upper
laminal cells of stem leaf; 12 – stem cross-section; 13-14 – perichaetial leaves; 15-16 – pseudoparaphyllia; 17 – upper
laminal cells of branch leaf; 18 – middle laminal cells of branch leaf; 19 – alar cells of branch leaf. Scale bars: 2 mm
– for 8; 1 mm – for 13-14; 0.5 mm – for 1-3, 5-7; 100 μm – for 4, 9-12, 15-19.
4
1
7
10
23
56
9
8
15
13 14 16 17
12
11
18
2 mm
100 μm
1 mm
0.5 mm
19
5. Stems regularly to irregularly pinnate-
branched; plants dioicous (never observed
with sporophytes in Altai) . . . . . . . . . . .. . 8
6. Plants small; leaves serrulate; alar cell
groups small and indistinct; pseudopara-
phyllia subulate numerous and conspicous
around branch initials; rare alpine epilith-
ic species . . . . . . . . H. recurvatum (p. 00)
6. Plants small to medium-sized; alar cell
groups rather distinct; pseudoparaphyl-
lia short and less numerous, not con-
spicous; common epixylic species . . . . . 7
7. Plants small-sized, leaves serrate to serrate;
alar cells rather homogeneous
. . . . . . . . . . . . . . . . . . . . . H. pallescens (p. 71)
7. Plants medium-sized, leaves serrulate; alar
cells heterogeneous . . . . H. fauriei (p. 80)
8. Leaves plicate, margins strongly revolute
. . . . . . . . . . . . . . . . . . . . H. revolutum (p. 74)
8. Leaves not plicate, margins erect or weak-
ly recurved below . . . . . . . . . . . . . . . . . . . 9
9. Stem leaves cordate-auriculate; large densely
and regularly pinnate alpine plants
. . . . . . . . . . . . . . . . . . . . H. procerrimum (p. 79)
9. Leaves otherwise . . . . . . . . . . . . . . . . . . . . . . . 10
10a. Large brownish alpine plants; leaves
markedly falcate; alar cells dark-brown,
thick-walled, forming an excavate group;
laminal cells thick-walled and porose,
elongate (8-10:1); pseudoparaphyllia
broadly ovate . . . H. bambergeri (p. 79)
10b. Medium-sized to large plants; leaves
weakly falcate to nearly straight; alar cells
form a ±well delimited square group; lam-
inal cells rarely porose, elongate (about
8-10:1); pseudoparaphyllia subfilamentose
to lanceolate . . H. cupressiforme (p. 76)
10c. Medium-sized plants; leaves concave,
±abruptly acuminate, usually falcate; alar
cells numerous, forming a less delimited
group; laminal cells not porose, relatively
short and wide (about 1:4-7); pseudopara-
phyllia ovate . . . . . . H. vaucheri (p. 75)
70 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 43. Hypnum pallescens (Hedw.) P. Beauv. (from Yailyu 600 m Ignatov 0/1715): 1, 6 – habit; 2 – capsule; 3 –
upper laminal cells of branch leaf; 4 – middle laminal cells of stem leaf; 5 – middle laminal cells of branch leaf; 7 –
upper laminal cells of stem leaf; 8-10 – stem leaves; 11-12 – pseudoparaphyllia; 13 – alar cells of stem leaf; 14-16 –
branch leaves; 17-18 – perichaetial leaves; 19 – alar cells of branch leaf; 20 – stem cross-section. Scale bars: 5 mm –
for 1; 2 mm – for 2, 6; 1 mm – for 8-10, 14-16, 17-18; 0.2 mm – for 11; 100 μm – for 3-5, 7 12-13, 19-20.
19
16
17 18
8
4
6
9
3
13
14
5
1
7
2
10
11
12
20
5 mm
2 mm 1 mm
100 μm
0.2 mm
15
Among the 11 species of Hypnum known
from Altai, the following species are discussed
worldwide in details by Ando: H. recurvatum
and H. pallescens – Ando, 1973; H. revolu-
tum – Ando, 1973, 1977; H. vaucheri – Ando,
1977; H. cupressiforme var. cupressiforme &
var. julaceum – Ando, 1989; var. subjulaceum
– Ando, 1990; var. filiforme – Ando, 1992; H.
bambergeri – Ando, 1996.
71
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1 2 3
4 5
Fig. 44. Hypnum pallescens (Hedw.) P. Beauv. (from Klyk Pass 700 m Ignatov 0/1718): 1 – peristome (290×); 2
– side view of exostome tooth in upper part (3000×); 3 – front view of exostome tooth in upper part (2500×); 4 – outer
surface of exostome tooth in lower part (5750×); 5 – outer surface of exostome tooth in transition zone (4300×).
Hypnum recurvatum (Lindb. & H. Arnell)
Kindb., Enum. Bryin. Exot. Suppl. 2: 100. 1891.
– Stereodon recurvatus Lindb. et H. Arnell,
Kongl. Sv. Vet. Akad. Handl. 23(10): 149. 1890.
Fig. 42
Plants small, in dense yellow-green tufts. Stem pros-
trate, to 2 cm long; epidermal cells non-differentiated;
central strand weak; regularly branched; branches 4
mm long. Pseudoparaphyllia subulate to filamentose,
numerous around branch primordia. Axillary hairs 3-
4-celled, 30-80 μm long, apical cell 20-40 × 6 μ m. Stem
leaves falcate, 0.6-0.8 × 0.2-0.3 mm, ovate-lanceolate;
margins recurved below, serrulate. Branch leaves 0.4-
0.6 × 0.1-0.2 mm. Laminal cells 35-50 × 4-5 (3-4/1-
1.5) μm. Alar cells few, weakly differentiated.
Autoicous. Sporophytes (old and imperfect)
present in a single Altaian collection (exostome de-
scription in part made using European material).
Perichaetial leaves erect, 2.2 × 0.8 mm, oblong-lan-
ceolate, shortly acuminate, plicate, costa reaching
about 2/3 the leaf length, margins entire. Seta 1.2
cm. Capsule ca. 1.2 mm long. Exostome teeth ca. 0.4
mm long, gradually tapered, nearly entire; striolate
below; in the middle densely papillose on many plates;
in the upper part very rarely papillose with small
irregularly arranged papillae, with prominent dor-
sal trabeculae and incrassate margin; ventral sur-
face and trabeculae smooth or with low irregular
papillae. Spores ca. 10 μm.
Distribution: Hypnum recurvatum has a scat-
tered distribution in mountains of Central and
Northern Europe, Caucasus, Siberia (both southern
and northern), Mongolia, Greenland, and North
America. In Altai it was found only once in the sub-
alpine zone, on rock outcrops.
Specimen examined: Bayas Lake 1860 m (0/1722).
Hypnum pallescens (Hedw.) P. Beauv.,
Prodr. Aetheogam. 67. 1805. – Leskea pallescens
Hedw., Sp. Musc. 219. t. 55, f. 1-6. 1801. – Hyp-
num reptile Michx., Fl. Bor. Am. 2: 315. 1803.
Figs. 43 & 44
Plants small, in dense dark-green to brownish
dull tufts. Stem prostrate, to 6 cm long; regularly
pinnate-branched; branches 4-5 mm long. Epider-
72 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 45. Hypnum revolutum (Mitt.) Lindb. (from Kukol 2400 m Ignatov 0/617): 1-3 – stem leaves; 4-5 – branch
leaves; 6 – habit; 7 – upper laminal cells of stem leaf; 8 – upper laminal cells of branch leaf; 9 – alar cells of stem leaf;
10 – alar cells of branch leaf; 11 – middle laminal cells of stem leaf; 12 – stem cross-section; 13-14 – pseudoparaphyl-
lia; 15 – middle laminal cells of branch leaf. Scale bars: 5 mm – for 6; 1 mm – for 1-5; 100 μm – for 7-15.
15
4
13
14
1
10
2
3
5
9
8
6
11
7
5 mm
1 mm
100 μm
12
mal cells not enlarged. Pseudoparaphyllia lanceolate
to subulate, often short. Axillary hairs 3-celled, ca. 70
μm, apical cell 30-40 × 5-6 μm. Stem leaves falcate-
secund, 1.0-1.6 × 0.4-0.6 mm, gradually tapered in
long narrow acumen, at base somewhat rounded;
margins distinctly to weakly serrulate throughout,
sometimes recurved below. Branch leaves 0.8-0.9 ×
0.2-0.3 mm. Laminal cells 35-55(-60) × 6 (5/1) μm.
In alar regions cells subquadrate, forming a large
indistinctly delimited group.
Autoicous. Inner perichaetial leaves 1.8-2.5 × 0.5-
0.6 mm, plicate, with indistinct double costa. Seta
0.8-1.2 cm. Capsule 1.3-1.5 mm long, arcuate, pale-
brown. Operculum long-rostrate. Exostome teeth 0.38
mm long, gradually tapered, serrulate in the upper
part; in the lower part striolate, with cristae some-
73
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 46. Hypnum vaucheri Lesq. (from Ulagan 1220 m Ignatov 36/236): 1-2 – habit; 3-5 – stem leaves; 6-7 – branch
leaves; 8 – stem cross-section; 9 – upper laminal cells of stem leaf; 10 – upper laminal cells of branch leaf; 11 – middle
laminal cells of branch leaf; 12 – middle laminal cells of stem leaf; 13-15 – pseudoparaphyllia; 16 – alar cells of branch
leaf; 17 – alar cells of stem leaf. Scale bars: 5 mm – for 1; 2 mm – for 2; 0.5 mm – for 3-7; 100 μm – for 8-17.
2 mm
0.5 mm
100 μm
15
4
13 14
1
7
10
2
35
9
12
8
11
17
6
16
5 mm
what wavy; in the middle densely papillose on many
plates; in the upper part with scattered low papil-
lae; dorsal lamelalle low to medium and more pap-
illose, margin non-incrassate; ventral surface and
trabeculae smooth or with rare low papillae. Endos-
tome to 0.4 mm long, basal membrane 0.15 mm high;
cilia nodose. Spores 12-15 μm.
Distribution: Widespread and common in
southern part of the boreal and northern parts of
the temperate zone throughout the Holarctic. In Altai
it is common on trunks, stumps and logs in forest
of at lower elevations, with very few findings in
the taiga zone.
Specimens examined: Ayukol 1000 m (0/1701); Ba-
yas Creek, in middle course 850 m (16/6); Bolshoi Shal-
tan Creek 540 m (0/1717); Gorno-Altaisk 400 m (35/
37); Kairu Creek, 8 km upstream 1000 m (15/116a; 15/
51); Karakol Lakes 1800 m (26/116); Kayakkatuyaryk-
skij Creek 1950 m (3/64); Kishte 440 m (Zolotukhin
11.VII.1988); Klyk Pass 700 m (0/1718); Kobukhtushka
440 m (Zolotukhina 13.VII.1988); Korbu 440 m (Zolo-
tukhin 20.X.1988); Srednij Shaltan Creek 640 m (0/
611); Tura Creek, in middle course 1300 m (28/26; 28/
48a; 28/50); Yailyu 450 m (0/1709), 480 m (0/628),
500 m (0/1716), 600 m (0/1715).
74 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 47. Hypnum cupressiforme Hedw. var. cupressiforme (a rather slender form, from Chulcha River, in middle course
1000 m Ignatov 9/156): 1-3 – stem leaves; 4-6 – branch leaves; 7 – pseudoparaphyllia; 8 – upper laminal cells of
stem leaf; 9 – upper laminal cells of branch leaves; 10 – middle laminal cells of stem leaf; 11 – middle laminal cells
of branch leaf; 12-13 – habit; 14 – alar cells of stem leaf; 15 – alar cells of branch leaf; 16 – stem cross-section. Scale
bars: 1 cm – for 13; 2 mm – for 12; 1 mm – for 1-6; 100 μm – for 7-11, 14-16.
12
15
14
25
11
7
6
8
9
34
10
13
1
16
2 mm
1 cm
1 mm
100 μm
Hypnum revolutum (Mitt.) Lindb., Oefv.
Foerh. Kongl. Sv. Vet.-Akad. 23: 542. (1866) 1867.
– Stereodon revolutus Mitt., J. Proc. Linn. Soc.,
Bot. Suppl. 1: 97. 1859. Fig. 45
Plants small to medium-sized, yellow-green to gold-
en-brownish mats. Stem prostrate to suberect, to 3 cm
long; epidermal cells not enlarged, but outer wall ±thin;
central strand weak; regularly pinnate-branched,
branches to 8 mm long. Pseudoparaphyllia short, ovate
to lanceolate. Axillary hairs 3-4-celled, 60-90 μm long,
apical cell 30-60 × 10 μm. Stem leaves crowded, falcate-
secund, 1.4-1.8 × 0.5-0.6 mm, more or less plicate; mar-
gins revolute nearly throughout, entire to slightly ser-
rulate above. Branch leaves 1.0-1.2 × 0.3-0.4. Laminal
cells 30-45 × 5-6 (4-5/1) μm. Alar subquadrate cells
form a rectangular group, ca. 10 cells long, 4-5 cells
wide. Dioicous. Sporophytes not found in Altai.
Distribution: This species is widespread in the
Arctic and in the alpine zone of most of the Holarc-
tic mountains. It has also several localities in the
Southern Hemisphere. In Altai it grows on rocks in
the alpine and subalpine belts.
Specimens examined: Kairu-Bazhi Peak 2300 m (13/
131); Kobiguayuk Creek 2300 m (0/199; 0/616); Kukol
2400 m (0/617); Tabozhok Peak 2350 m (31/320), 2700
m (31/308; 31/309), 2750 m (31/311); Yakhansoru Lake
2000 m (Zolotukhin 26.VI.1990).
75
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 48. Hypnum cupressiforme var. filiforme Brid. (from Kayakkatuyarykskij Creek 1850 m Ignatov 8/156): 1-2 –
habit; 3-5 – stem leaves; 6-8 – branch leaves; 9 – stem cross-section; 10 – middle laminal cells of branch leaf; 11 –
alar cells of stem leaf; 12 – pseudoparaphyllia; 3 – upper laminal cells of branch leaf; 14 – upper laminal cells of stem
leaf; 15 – alar cells of branch leaf; 16 – middle laminal cells of branch leaf. Scale bars: 1 cm – for 1; 2 mm – for 2;
1 mm – for 3-8; 100 μm – for 9-16.
16
11
8
12
2
1
14
7
9
4
6
5
15
10
3
13
1 cm 2 mm
1 mm
100 μm
Hypnum vaucheri Lesq., Mém. Soc. Sci. Nat.
Neuchatel 3(3): 48. 1846. Fig. 46
Plants small to medium-sized, in compact green,
yellow-green or yellow-brown mats. Stem prostrate
to erect-ascending (often branches are erect from
prostrate stem); ca. 2 cm long; regularly branched,
branches to 6 mm long. Pseudoparaphyllia semior-
bicular to rounded-rectangular. Axillary hairs 3-celled,
40-60 μm, apical cell 20-30 × 9-10 μm. Stem leaves
falcate, 0.9-1.3 × 0.4-0.6 mm, ovate, concave; margins
plane, involute in the middle, subentire. Branch leaves
0.9-1.0 × 0.3-0.4 mm. Laminal cells 30-35(-40) × 7-8
(5-6/2). Alar cells subquadrate, forming an exten-
sive indistinctly delimited group, 10-15 cells along
the margin. Dioicous. Sporophyte not seen in Altai.
Hypnum vaucheri is similar to some forms of H.
cupressiforme, especially its var. julaceum, and some
authors even consider these speceis to be conspecific
(cf. Crum & Anderson, 1981). Sometimes the shape of
paraphyllia (subfilamentose to lanceolate in H. cu-
pressiforme/ broadly ovate in H. vaucheri) needs
to be examined, though in most cases shorter lami-
nal cells and numerous angular cells are enough to
recognize this species.
Distribution: This species has a wide distribu-
tion throughout the Holarctic, obviously preferring
more xeric regions within the Arctic, forest- and steppe
zones. In Altai it grows at all elevations, on soil
and rocks, sometimes in very dry steppes.
Specimens examined: Azhi Cape 440 m (21/34); Ba-
yas Lake 1850 m (0/614); Berekhtuyaryk 1600 m (0/
612); Chainary Creek, upper course 1800 m (34/199);
Chemal 400 m (29/39; 29/40); Chemal, 10 km upstream
along Katun 450 m (34/145; 34/170); Derbogach 450 m
(0/33); Gorno-Altaisk (SE edge) 600 m (23/16); Kairu
Creek, 8 km upstream 1000 m (15/38); Kairu-Bazhi Peak
2050 m (13/171), 2100 m (13/81); Kobukhta 550 m (0/
621), 600 m (0/226); Kokorya Creek 2400 m (32/51);
Kukol 1800 m (0/613; 0/624), 2000 m (0/620); Malyi
Yaloman 900 m (25/154; 25/30), 1050 m (25/136; 25/
82), 1200 m (25/131); Malyi Yaloman Creek, 2 km up-
76 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 49. Hypnum cupressiforme var. julaceum Brid. (from Kayakkatuyarykskij Creek 1930 m Ignatov 3/189): 1-2 –
habit; 3-5 – stem leaves; 6-8 – branch leaves; 9, 13 – pseudoparaphyllia; 10 – upper laminal cells of stem leaf; 11 –
middle laminal cells of stem leaf; 12 – alar cells of stem leaf; 14 – upper laminal cells of branch leaf; 15 – middle
laminal cells of branch leaf; 16 – alar cells of branch leaf. Scale bars: 5 mm – for 1; 2 mm – for 2; 1 mm – for 3-8;
100 μm – for 9-16.
15
5
16
8
4
10
7
6
12
11
9
13
2
1
14
3
2 mm
1 mm
100 μm
5 mm
stream 850 m (25/168), 1100 m (25/169); Tabozhok Creek,
12 km upstream 2200 m (30/132; 30/134; 30/139; 30/
141); Tabozhok Creek, 8 km upstream 2050 m (30/133;
30/146; 30/147), 2250 m (30/136; 30/137), 2500 m (30/
135; 30/144); Tabozhok Peak 2350 m (31/306; 31/313;
31/315; 31/321; 31/322), 2450 m (31/307); Tokpak
Creek, upper course 2400 m (36/386); Ulagan 1220 m
(36/236); Ust-Sema 350 m (24/77); Yailyu 450 m (0/
622); Yakhansoru Lake 1870 m (Zolotukhin 28.VI.1990);
Yazula 1600 m (0/618).
Hypnum cupressiforme Hedw., Sp. Musc.
291. 1801. Figs. 47–51
Plants medium-sized to rather robust, green, yel-
low to brownish. Stem prostrate to erect, complanate
to subjulaceous, 3-4(8) cm long; epidermal cells not
differentiated; regularly pinnate-branched to near-
ly unbranched; branches to 7(12) mm long. Pseudo-
paraphyllia subfilamentose to lanceolate, sometimes
fused at base and bi- or trilobed. Axillary hairs 4-5-
celled, 100-150 μm long, apical cell ca. 50 × 11 μm.
Leaves falcate to nearly straight (1.1-)1.4-1.8(-2.0)
× (0.4-).5-0.6(-0.7) mm, ovate-lanceolate; margins
serrulate above, often recurved below. Laminal cells
40-75 × 5-7 (4-5/1.5-2) μm. Alar subquadrate cells
numerous, ±distinctly delimited.
Dioicous. No sporophytes observed in Altai, or in
other inland regions of Russia. The following de-
scription of sporophytes was made from European
specimens. Perichaetial leaves erect, plane. Seta ca. 2
cm. Capsule ca. 2 mm long. Operculum rostrate. An-
nulus of 2-3 row of cells, deciduous. Exostome teeth
ca. 0.5 mm long, gradually tapered, nearly entire;
perfectly striolate below; in the middle densely pap-
illose on many plates; in the upper part with low
medium papillae in longitudinal rows; dorsal trabe-
culae low-papillose, with non-incrassate margin; ven-
tral surface and trabeculae smooth or with rare low
papillae. Endostome as long as the teeth, basal mem-
brane about 0.2 mm high; segments perforate; cilia
nodose. Spores ca. 15 μm.
77
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 50. Hypnum cupressiforme var. subjulaceum Mol. (from Bogoyash Creek, upper course 2550 m Ignatov 36/255):
1-4 – leaves; 5 – middle laminal cells; 6-7 – alar cells; 8 – upper laminal cells; 9 – pseudoparaphyllia; 10-11 – habit.
Scale bars: 1 cm – for 10; 2 mm – for 11; 1 mm – for 1-4; 100 μm – for 5-9.
8
4
10
6
11
9
3
5
1
7
2
1 mm
100 μm
1 cm
2 mm
In Altai this species is enormously polymor-
phous, represented by var. cupressiforme (com-
mon in the forest, rarely in the subalpine zones,
typically on trunks and in diffuse shade); var.
filiforme (rather rare, nearly at the same eleva-
tions, but usually on deeply shaded rocks); var.
julaceum (rare at about the tree-line), and var.
subjulaceum (usually in the subalpine and al-
pine zones, but with a number of localities in
the forest zone; usually on well exposed rocks).
The latter variety is the most peculiar and some
authors consider it as a separate species. Hyp-
num cupressiforme var. subjulaceum has a ten-
dency to show complanate foliage (superficial-
ly resembling Callicladium). Alar cells are typ-
ically strongly colored, thick-walled, porose, and
form a well-delimited concave group. However
in many collections of var. subjulaceums some
marginal or submarginal alar cells became larger,
more transparent, with rather thin, non-porose
walls. A key to the 9 varieties recognized with-
in Hypnum cupressiforme was published by
Ando (1993).
var. cupressiforme Fig. 47
Plants ±regularly and often densely pinnate-
branched, usually green. Stem leaves falcate-secund,
1.2-2.0 × 0.4-0.6 mm, subentire to slightly serrulate.
Branch leaves 1.0-1.5 × 0.3-0.4 mm. Laminal cells
40-70 × 6-7(4-5/2) μm, in the alar region subquad-
rate cells opaque, thick-walled, forming a distinctly
delimited group.
Specimens examined: Akkurum, 3 km downstream by
Chulyshman: 550 m (Zolotukhin 22.IX.1989); Ayukol:
1100 m (0/1694); Bayas Creek, at mouth: 500 m (0/
1699); Chiri Creek, 0.5 km upstream: 450 m (17/104;
17/71); Chulcha River, in middle course: 1000 m (9/
156); 1100 m (9/52); Kairu Creek, 8 km upstream: 1000
m (15/69); Kairu-Bazhi Peak: 2100 m (13/43); Karagai:
440 m (0/271); Kayakkatuyarykskij Creek: 1930 m (3/
223); 2100 m (7/121); Korbu: 440 m (0/1696); Kukol:
1750 m (0/1708); Kyga River, 2 km upstream: 600 m
(0/1700); Srednij Shaltan Creek: 730 m (0/1697);
Tura Creek, in middle course: 1300 m (28/10; 28/18);
Yailyu: 440 m (0/1693; 0/1698); 450 m (0/632);
Yakhansoru Lake: 1850 m (Zolotukhin 28.VI.1990).
var. filiforme Brid., Musc. Rec. 2(2): 138.
1801. Fig. 48
Plants irregularly to regularly, but not densely
pinnate-branched, usually green. Stem leaves nearly
straight, 1.0-1.4 × 0.3-0.4 mm, less serrulate than type
variety to nearly entire. Branch leaves 1.0-1.2 × 0.25-
0.3 mm. Laminal cells 40-75 × 5(3/2) μm, in the alar
region subquadrate cells opaque, thick-walled, form-
ing a distinctly delimited group.
78 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1
2
3 4
Fig. 51. Hypnum cupressiforme Hedw. var. cupressiforme (from Finland, Brotherus, 291C): 1 – peristome (210×); 2
– outer surface of exostome tooth in upper part (3500×); 3 – outer surface of exostome tooth in lower part (5000×); 4
– outer surface of exostome tooth in transition zone (4000×).
Specimens examined: Chulcha River, in middle course:
1200 m (9/24); Kamga Creek, 11 km upstream: 560 m
(0/1524); Kayakkatuyarykskij Creek: 1850 m (8/156).
var. julaceum Brid., Spec. Musc. 2: 216. 1812.
Fig. 49
Plants irregularly pinnate-branched, green in
young parts to yellow brown, julaceous. Stem leaves
1.1-1.2 × 0.4-0.5 mm, from the ovate base abruptly
and shortly erect-acuminate, subentire. Branch leaves
somewhat larger, 1.3 × 0.6-0.7 mm. Laminal cells 40-
50(-60) μm.
Specimens examined: Kayakkatuyarykskij Creek: 1930
m (3/189)
var. subjulaceum Mol., Ber. Naturh. Ver. Augs-
burg. 18: 183. 1865. Fig. 50
Plants irregularly to regularly, but not densely
pinnate-branched, usually yellow-brown (green in
the uppermost portion). Stem leaves nearly straight,
1.3-1.5(-2.0) × 0.4-0.5(0.7) mm, serrulate to suben-
tire. Branch leaves slightly differentiated. Laminal
cells 45-75 × 6(4/2) μm, thick-walled and at places
distinctly porose; in the alar region sunquadrate cells
opaque, thick-walled and dark brown, forming a
distinctly delimited excavate group; sometimes alar
cells partly inflated.
Specimens examined: Ayulyuyuzyuk Creek 2050 m (0/
165), 2150 m (0/128; 0/377); Bogoyash Creek, upper
course 2550 m (36/255); Chulcha River, in middle course
850 m (Zolotukhin 18.IX.1989), 960 m (9/92); Kairu-
Bazhi Peak 2300 m (13/130); Kayakkatuyarykskij Creek
1760 m (8/43), 1850 m (8/288), 1970 m (7/125), 2000 m
(3/272); Kobiguayuk Creek 2300 m (0/203); Kukol 1800
m (0/1524; 0/1731), 1900 m (0/1523); Ok-Porok Creek,
at mouth 440 m (0/634a); Srednij Shaltan Creek 740 m
(0/1520); Tabozhok Peak 2300 m (31/316), 2700 m (30/
142); Yailyu 450 m (0/1523); Yakhansoru Lake 1870 m
(Zolotukhin 28.VI.1990).
79
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 52. Hypnum bambergeri Schimp. (from Tabozhok Peak 2800 m Ignatov 0/2030): 1,8 – habit; 2-4 – leaves;
5 – middle laminal cells; 6 – pseudoparaphyllium; 7 – stem cross-section; 9 – upper laminal cells; 10 – alar cells.
Scale bars: 1 cm – for 8; 5 mm – for 1; 1 mm – for 2-4; 100 μm – for 5-7, 9-10.
1 mm
1 cm
100 μm
4
1710
2
3
9
8
6
5 mm
5
Transitions between var. cupressiforme and
var. subjulaceum are found in specimens:
Bogoyash Creek, upper course 2400 m (36/245); Chain-
ary Creek, upper course: 1800 m (34/200); Chiri Creek,
0.5 km upstream: 450 m (17/74); Tabozhok Peak 2700
m (31/312).
Transitions between var. subjulaceum and
var. filiforme in specimen:
Chemal Creek, 3 km upstream 450 m (34/32).
Hypnum bambergeri Schimp., Syn. Musc.
Eur., ed. 2, 2: 698. 1860. Fig. 52
Plants medium-sized to large, yellow-green in young
parts, soon becoming brown. Stem prostrate to ascend-
ing, 4 cm long; without hyalodermis and with weak
central strand; branches few, to 8 mm long. Pseudopara-
phyllia broadly-ovate to broadly triangular. Axillary hairs
3-4-celled, 120-140 μm long, apical cell 40-65 × 10 μm.
Stem leaves crowded, falcate to circinate-secund, 1.5-1.8
× 0.5-0.6 mm; margins plane, entire. Branch leaves 1.3-
1.4 × 0.3-0.4 mm. Laminal cells 40-65 × 6-7 (3.5-5/2.0-
2.5) μm, thick-walled, porose. Alar cells subquadrate to
rectangular, thick-walled, forming a distinct brown ex-
cavate group. Dioicous. Sporophytes not seen in Altai.
Distribution: Hypnum bambergeri occurs in the
Arctic and in high mountains of Europe, Azores, South
Siberia, in North America to British Columbia in the
west, and to Newfoundland and Quebec in the east.
In Altai it was found in few places at or above the
tree-line, on rocks or in wet tundras.
Specimens examined: Tabozhok Peak 2550 m (0/2063),
2800 m (0/2030; 0/2036).
Hypnum procerrimum Mol., Flora 49: 458.
1866. – Pseudostereodon procerrimum (Mol.)
Fleisch. in Broth., Nat. Pfl. (ed. 2) 11: 455. 1925.
– Ctenidium procerrimum (Mol.) Lindb., Bot.
Not. 1882: 26. 1882. Fig. 53
Plants medium-sized, in dense golden-brownish
tufts. Stem prostrate, 2-3 cm long; without hyaloder-
mis, with weak central strand; ±regularly but rarely
branched, branches 4 mm long. Pseudoparaphyllia
broadly-triangular to transversely ovate. Axillary
hairs 4-9-celled, all colored, 140-200 μm long, apical
cell 50-60 × 10-12 μm. Stem leaves strongly falcate,
2.5-2.8 × 0.6-0.7 mm, cordate-auriculate at base, pli-
cate, margin plane. Branch leaves 1.4-1.5 × 0.3-0.4
mm. Laminal cells 40-60 × 6 (3.5/2.5) μm, thick-
walled, porose. Alar cells isodiametric, thick-walled,
in a large indistinctly delimited group, lowermost
cells sometimes inflated. Dioicous. Sporophytes un-
known in this species.
This species has leaves with a peculiar cor-
date-auriculate base and weakly differentiated
alar cells and for these reasons is placed some-
times in Ctenidium (cf. Corley & al., 1981;
Nyholm, 1965). However other characters, such
as entire strongly falcate leaves with smooth
(or nearly so) laminal cells, suggest its more
close relation to Hypnum, rather than to Ctenid-
ium. The segregation of this species in the mo-
notypic genus Pseudostereodon is possible in
case of further splitting of Hypnum.
80 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 53. Hypnum procerrimum Mol. (from Bogoyash Creek, upper course 2400 m Ignatov 36/66): 1-2 – stem leaves;
3, 7 – pseudoparaphyllia; 4-6 – branch leaves; 8 – habit; 9 – upper laminal cells of stem leaf; 10 – upper laminal
cells of branch leaf; 11 – middle laminal cells of stem leaf; 12 – middle laminal cells of branch leaf; 13 – alar cells
of branch leaf; 14 – alar cells of stem leaf; 15 – stem cross-section. Scale bars: 5 mm – for 8; 1 mm – for 1-2, 4-6; 100
μm – for 3, 7, 9-15.
15
2
5
8
9
11
12
13
7
6
3
1
4
14
10
5 mm
100 μm
1 mm
Distribution: Hypnum procerrimum is more or
less widely distributed in the Arctic and in high
mountains of Central and South Europe, Caucasus,
Siberia, Middle Asia, Mongolia, China (southward
to Yunnan), and British Columbia and Alberta in
North America. In Altai this species occurs in xeric
tundras of South-East Altai, on rocks or soil.
Specimens examined: Ak-Turu Creek, upper course 2600
m (Bardunov 18.VII.1966 IRK!); Bogoyash Creek, upper
course 2400 m (36/66), 2450 m (36/252); Kokorya Creek
2400 m (32/50); Tabozhok Peak 2350 m (31/314), 2400
m (31/331); Tutuoyuk Creek, upper course 2500 m (Zolo-
tukhin & al. 15.VII.1990).
Hypnum fauriei Card., Beih. Bot. Centralbl.
17: 41. 26. 1904. Figs. 54 & 55
Plants medium-sized, in soft green to yellowish-
green tufts. Stem prostrate or ascending when grow-
ing in dense tufts, to 5 cm long; regularly densely
branching, with branches ca. 7 mm long. Epidermal
cells indistinct to enlarged at places. Pseudopara-
phyllia lanceolate-subulate, sometimes fused at base.
Axillary hairs 3-celled, 70-120 μm long, apical cell
40-75 × 8 μm. Stem leaves strongly falcate, 1.7-2.1 ×
0.5-0.7 mm, gradually tapered in long narrow acu-
men; margins serrulate above. Branch leaves 1.1-1.4
× 0.4 mm. Laminal cells 50-80 × 4(3/1) μm. In alar
regions cells subquadrate to rectangular, some low-
ermost – enlarged and thin-walled, forming a rath-
er distinctly (in stem leaves) or indistinctly (in
branch leaves) delimited group.
Autoicous. Sporophytes frequent. Perichaetial
leaves 1.8-2.5 × 0.6 mm, entire to serrulate, plicate, with
indistinct double costa. Seta 2-3 cm. Capsule 15-20(25)
mm long, arcuate, slightly furrowed when dry. Oper-
81
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 54. Hypnum fauriei Card. (from Yurga 450 m Ignatov 0/1713): 1,3 – habit; 2 – capsule; 4-7 – pseudopara-
phyllia; 8 – stem cross-section; 9-10 – perichaetial leaves; 11-13 – stem leaves; 14 – alar cells of stem leaf; 15-17 –
branch leaves; 18 – middle laminal cells of stem leaf; 19 – middle laminal cells of branch leaf; 20 – upper laminal
cells of stem leaf; 21 – upper laminal cells of branch leaf; 22 – alar cells of branch leaf. Scale bars: 1 cm – for 1; 2
mm – for 2-3; 1 mm – for 9-13, 15-17; 100 μm – for 4-8, 14, 18-22.
20 21 22
13
19
68
11
4
12
7
3
14
15
9
1
5
2
18
17
16
100 μm
1 cm
2 mm
10
1 mm
culum conic, shortly acuminate. Exostome teeth 0.55
mm long, gradually tapered, distinctly serrate in the
upper third; imperfectly striolate below; in the mid-
dle densely papillose on many plates; in the upper
part with rare minute papillae arranged in indistinct
rows, and high-conic papillae along relatively low
dorsal trabeculae and non-incrassate margin; ventral
surface and trabeculae with dense conic-spinulose
papillae. Endostome 0.65 mm long, basal membrane
0.24 mm high; cilia nodose. Spores 16-20 μm.
Hypnum fauriei in Altai is distinct due to
the medium size of plants with dense regularly
pinnate branching. This species is close to H.
fertile Sendtn., but differs from it in: (1) stem
leaves wider lanceolate, 1,4-2.2 × 0.45-0.75 mm
vs. 1.3-2.2 × 0.4-0.5 mm; (2) median laminal
cells shorter, (40-)50-70(-80) μm long vs. (50-
)60-80(-90) μm; (3) capsule longer, 2-3 mm long
(urn) vs. 1.5-2.5 mm; (4) spores larger, 15-22(-
24) μm in diameter vs. (10-)13-16
(-20) μm. Previous records of H. fertile from
82 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1
2
4
5
3
Fig. 55. Hypnum fauriei Card. (from Karagai 440 m
Ignatov 0/1710): 1 – peristome (190×); 2 – side view
of exostome tooth in upper part, showing echinate-papil-
lose inner trabeculae (5000×); 3 – front view of exos-
tome tooth in upper part (4000×); 4 – outer surface of
exostome tooth in transition zone and below (4400×); 5
– outer surface of exostome tooth in lower part (4400×).
83
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 56. Hypnum hamulosum B.S.G. (from Kobiguayuk Creek 2850 m Ignatov 0/223): 1-2 – habit; 3-4 – stem
leaves; 5-6 – branch leaves; 7 – middle laminal cells of stem leaf; 8 – alar cells of stem leaf; 9 – pseudoparaphyllium;
10 – stem cross-section; 11 – upper laminal cells of stem leaf; 12 – upper laminal cells of branch leaf; 13 – middle
laminal cells of branch leaf; 14 – alar cells of branch leaf. Scale bars: 3 mm – for 1; 2 mm – for 2; 0.5 mm – for 3-
6; 100 μm – for 7-14.
3 mm
0.5 mm
2 mm
100 μm
8
4
6
11
9
3
5
17
2
10
13 14
12
Altai (Ignatov, 1990) and other parts of Sibe-
ria (Brotherus, 1925) are based on misidentifi-
cations of H. fauriei.
Distribution: Hypnum fauriei occurs in Japan,
Korea, China, Russian Far East and South Siberia,
westward to Altai. In Altai it grows at lower eleva-
tions in the areas with especially mild climate
(around Teletzkoye Lake and Katun River valley
near Chemal). All collections were made from rela-
tively dry rotten logs in partial shade.
Specimens examined: Chemal Creek, 3 km upstream
450 m (34/231); Chichelgan Range 800 m (0/1725);
Kairu Creek, 8 km upstream 1000 m (15/116; 15/144);
Karagai 440 m (0/1710); Kobukhta 450 m (0/1712);
Yailyu 480 m (0/631); Yurga 450 m (0/1713).
Hypnum hamulosum B.S.G., Bryol. Eur. 6:
96. pl. 590. 1854. Fig. 56
Plants small for the genus, in soft green to yel-
lowish-brown dense tufts. Stem prostrate to ascend-
ing, to 2 cm long; with enlarged epidermal cells;
irregularly to ±regularly branching, with branches
ca. 4 mm long. Pseudoparaphyllia lanceolate to sub-
ulate. Axillary hairs 3-celled, 60-70 μm long, apical
cell 35-40 × 8 μm. Stem leaves strongly falcate, 1.0-
1.2 × 0.3-0.4 mm, ±plicate; margins recurved below,
serrulate above. Branch leaves 0.7-0.9 × 0.2-0.3 mm.
Laminal cells 40-50 × 4-5 (3-4/1) μm. In alar re-
gions few cells subquadrate, forming an indistinct
group, sometimes 1-2 corner cells enlarged.
Dioicous. Sporophytes were observed in Altai in
only one collection (deep wet canyon in mild cli-
mate). Perichaetial leaves 2.0 × 0.4 mm, plicate, coarse-
ly serrate above, serrulate below. Seta 1.5 cm. Cap-
sule 2 mm long. Exostome teeth 0.35 mm long, grad-
ually tapered, serrate; striolate below; in the middle
densely papillose on many plates; in the upper part
moderately papillose with low irregularly arranged
papillae, with low dorsal trabeculae and non-incras-
sate margin; ventral surface and trabeculae smooth.
Endostome normal for the genus, cilia nodose or some-
what appendiculate. Spores 11-14 μm.
Hypnum hamulosum is easy to know by
small plants with leaves strongly falcate, wide-
ly spreading from the stem or branches imme-
84 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 57. Hypnum plicatulum (Lindb.) Jaeg. (from Kayakkatuyarykskij Creek 2100 m Ignatov 7/120): 1 – habit; 2-
4 – stem leaves; 5 – middle laminal cells of stem leaf; 6-8 – branch leaves; 9-10, 14 – pseudoparaphyllia; 11 – alar
cells of stem leaf; 12 – upper laminal cells of stem leaf; 13 – stem cross-section; 15 – upper laminal cells of branch
leaf; 16 – middle laminal cells of branch leaf; 17 – alar cells of branch leaf. Scale bars: 2 mm – for 1; 1 mm – for 2-
4, 6-8; 100 μm – for 5, 9-17.
1
2
4
5
3
7
6
11
13
10
9
15
14
16 17
1 mm
8
100 μm
12
2 mm
diately from the insertion (while in other small
Hypnum species occurring in Altai the leaf base
is ±appressed to the stem). A key for the spe-
cies of the section Hamulosa has been published
by Ando (1994).
Distribution: This species has a scattered distri-
bution in the Holarctic, mostly in northern and
mountain areas. It is rather common in Siberian
mountains. In Altai it grows usually on wet and
partially shaded rock outcrops (especially close to
waterfalls), in all the altitudinal zones.
Specimens examined: Bolshoye Istyube Creek 470 m
(Zolotukhin 5.VI.1986; 0/629; 18/127; 18/14); Chich-
elgan Cape 440 m (Zolotukhina 12.VII.1988); Chiri Creek,
0.5 km upstream 450 m (17/10); Kairu Creek, 16 km up-
stream 1400 m (14/18); Kairu Creek, 8 km upstream 1000
m (15/141); Kairu-Bazhi Peak 1700 m (13/92); Kamga
Creek, 10 km upstream 500 m (0/104), 560 m (0/1704),
580 m (0/1721); Kamga Creek, 14 km upstream 660 m
(0/1719); Karakol Lakes 1950 m (28/53); Kayakkatu-
yarykskij Creek 2050 m (3/59; 5/4); Kobiguayuk Creek
2850 m (0/223); Korbu 440 m (Zolotukhin 20.X.1988);
Ok-Porok Creek, 1.5 km upstream 470 m (2/27); Srednij
Shaltan Creek 670 m (0/1702); Yailyu 450 m (0/1703).
85
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 58. Hypnum pratense J. Koch ex Spruce (from Gorno-Altaisk 400 m Ignatov 35/49): 1-2 – stem leaves; 3-4 –
branch leaves; 5 – upper stem; 6 – habit; 7 – upper lamina cells of stem leaf; 8 – middle lamina cells of stem leaf;
9 – alar cells of branch leaf. 10 – proximal branch leaf; 11-13 – pseudoparaphyllia; 14 – stem cross-section. Scale bars:
5 mm – for 6; 2 mm – for 5; 1 mm – for 1-4; 100 μm – for – 7-14.
1
234
5
6
7
8
9
10 11 12 13 14
100 μm
2 mm
1 mm
5 mm
Hypnum plicatulum (Lindb.) Jaeg., Ber. St.
Gall. Naturw. Ges. 1877-78: 316. 1880. – Stere-
odon plicatulus Lindb., Acta Soc. Sci. Fenn. 10:
251. 1872. Fig. 57
Plants rather small, in yellow-green tufts. Stem
prostrate, to 1.5 cm long; epidermal cells differenti-
ated, but at places indistinctly so; regularly pinnate-
branched (young branches elongate evenly, so shoot
ends have usually characteristically triangular out-
line), branches to 4 mm. Pseudoparaphyllia lanceolate
to subulate. Axillary hairs 3-celled, 90 μm long, api-
cal cell ca. 60 × 10 μm. Stem leaves falcate 1.0-1.5 ×
0.5-0.8 mm, triangular-lanceolate, rounded to cor-
date base, acuminate; margins subentire, plane. Branch
leaves 0.9-1.5 × 0.4-0.5 mm. Laminal cells 55-70 × 4
(3/1) μm, in alar regions cells subquadrate, thin-
walled, forming a small indistinct group. Dioicous.
Sporophytes not seen in Altai.
Distribution: Hypnum plicatulum occurs most-
ly in Asia: Ural Mts. (from the Arctic to southern
Urals), most of the Asian Arctic territories, moun-
tains of South Siberia, Russian Far East, Mongolia,
Korea and Japan. Outside Asia it is known from the
Arctic in European Russia, North America and Green-
land. In Altai this species grows in moist rocks (sim-
ilar to H. hamulosum), in all the altitudinal zones
(except the upper alpine), obviously preferring the
subalpine belt. Schofield (1992) mapped the distri-
bution of this species in North America.
Specimens examined: Ayukol 1000 m (0/630); Bayas
Lake 1750 m (0/1720; 0/610); Karakol Lakes 1900 m
(28/157; 28/54); Kayakkatuyarykskij Creek 1950 m (3/
53), 2100 m (7/120); Kishte 440 m (Zolotukhin
11.VII.1988); Kobiguayuk Creek 2150 m (0/318);
Uzunkel Lake 2050 m (Zolotukhin 23.VI.1990).
Hypnum pratense J. Koch ex Spruce, Lon-
don J. Bot. 4:177. 1845. – Breidleria pratensis
(J. Koch ex Spruce) Loeske, Stud. Morph. Syst.
Laubm. 172. 1910. Figs. 58 & 59
Plants medium-sized to robust, in soft glossy light
yellow-green tufts. Stem prostrate to ascending, dense-
ly complanate, to 4 cm long; with hyalodermis and
86 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1
2
3
Fig. 59. Hypnum pratense J. Koch ex Spruce (from
Central Russia, Yaroslavl Province, Zickendrath,
31.V.1891, ex MW): 1 – peristome (110×); 2 – outer
surface of exostome tooth in upper part (4400×); 3 –
inner surface of exostome tooth in upper part (3600×).
central strand; irregularly-branched. Pseudoparaphyl-
lia broadly to narrowly lanceolate. Axillary hairs 4-
5-celled, 80-120 μm long, apical cell 50-60 × 8 μm.
Stem leaves straight or weakly falcate, 1.8-2.3 × 0.7-
0.9 mm, ovate-lanceolate, gradually acuminate, ser-
rulate above. Branch leaves 1.3-1.4 × 0.4-0.5 mm.
Laminal cells 60-85 × 4-5(3-4/1) μm, at base porose,
in alar regions subquadrate, thin-walled, forming a
indistinctly delimited group.
Dioicous. Sporophyte not found in Altai, and very
rare else where in the range. The following description
of one specimen from Central Russia may be useful.
Perichaetial leaves plicate. Seta ca. 3 cm, Capsule 3 mm
long. Exostome teeth ca. 0.4 mm long, abruptly tapered,
serrulate in the distal third; striolate below; in the middle
densely papillose; in the upper part with rare, irregu-
larly arranged papillae; dorsal trabeculae and incras-
sate margins prominent, nearly smooth; ventral trabe-
culae smooth. Endostome cilia nodose.
The genus Breidleria is segregated sometimes
from Hypnum for the species having large plants
and large epidermal cells. Several species were
placed in this genus (see the review by Hedenäs,
1992), but most manuals recognize it for two wide-
spread species Hypnum (Breidleria) pratense and
Hypnum (Breidleria) lindbergii. Hedenäs (1992)
found the latter species to be more close to Calli-
ergonella cuspidata, and transferred H. lindbergii
to Calliergonella. Our observations on exostome
ornamentation increase the number of differences
between Hypnum pratense and Calliergonella
lindbergii. However Calliergonella cuspidata
and C. lindbergii have numerous differences
which will be discussed elsewhere. In Breidleria
Hedenäs left Hypnum pratense and close East-
Asian H. erectiusculum Sull. et Lesq. However,
Hypnum at present is too heterogeneous to con-
sider Breidleria (as well as Pseudostereodon) to
be a good segregate, though further splitting of
Hypnum may resurrect Breidleria.
Distribution: The species is known from most bo-
real and north-temperate areas of the Northern Hemi-
sphere, but in most of the areas it is a rare moss. In Altai
it was found once in a mesic Betula forest, on soil.
Specimen examined: Gorno-Altaisk 400 m (35/49).
DOUBTFUL RECORD
Hypnum callichroum Brid. was reported for
Altai by Krylov (1924). The specimen was not found,
but all so-named collections from South Siberia
were found to be H. plicatulum, so we doubt that
H. callichroum occurs in South Siberia.
Callicladium Crum
This genus was established by Crum (1971)
to accommodate one widespread north-temperate
and hemiboreal species, Callicladium haldani-
anum. For a long time it was placed in Hetero-
phyllium, due to the somewhat complanate habit,
conspicuous groups of thin-walled alar cells, and
autoicous sexual condition. However this posi-
tion was found not appropriate because of thin-
walled alar cells. Essential differences from Hyp-
87
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 60. Callicladium haldanianum (Grev.) Crum (from Klyk Pass 700 m Ignatov 0/636): 1 – habit; 2-4 – stem
leaves; 5, 12 – shoot ends, showing variations; 6, 13-14 – pseudoparaphyllia; 7 – stem cross-section; 8-10 – branch
leaves; 11 – capsule; 15-16 – upper laminal cells of stem leaves; 17-18 – middle laminal cells of stem leaves; 19-21
– perichaetial leaves; 22 – alar cells of perichaetial leaf; 23-24 – alar cells of stem leaves. Scale bars: 1 cm – for 1;
2 mm – for 5, 11-12; 1 mm – for 2-4, 8-10, 19-21; 100 μm – for 6-7, 13-18, 22-24.
7
1
234
5
6
8910
11
12
13
14
15
100 μm
21
22 23 24
16
18
19 20
2 mm
1 cm
1 mm
17
num include subjulaceous branches and ±straight
leaves with entire margins. However some forms
of H. cupressiforme (especially var. subjulaceum
and transitional forms between var. subjulaceum
and var. filiforme) have a general appearance very
close to that of Callicladium and they are easy to
confuse. Their differences include sexual condi-
tions (autoicous in Callicladium, dioicous in H.
cupressiforme) and thin-walled alar cells in Cal-
licladium (porose in most leaves H. cupressiforme
var. subjulaceum, though in some leaves the alar
groups consist of mostly translucent and not
88 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 2
43
Fig. 61. Callicladium haldanianum (Grev.) Crum (from Yailyu, 450 m, Ignatov, 0/1521): 1 – peristome (200×); 2
– side view of exostome tooth in upper part (5300×); 3 – outer surface of exostome tooth in lower part (5600×); 4 –
outer surface of exostome tooth in transition zone (4400×).
especially porose cells). Similarity with the Hyp-
num cupressiforme-group (Hypnum sect. Hypnum)
includes also non-plicate perichaetial leaves, which
in all other sections of Hypnum are plicate. There-
fore the generic segregation of Callicaldium poses
a question about the further splitting of Hypnum.
Callicladium haldanianum (Grev.) Crum,
Bryologist 74: 167. 1971. – Hypnum haldani-
anum Grev., Ann. Lyc. Nat. Hist. NY 1: 275. 1825.
– Heterophyllium haldanianum (Grev.) Kindb.,
Canad. Rec. Sci. 6: 72. 1894. Figs. 60 & 61
Plants large, in green to yellow-green glossy tufts.
Stem prostrate, to 6 cm long; without hyalodermis;
subjulaceous except distinctly complanate and always
slightly arcuate-ascending tips; ±regularly pinnate-
branched, branches 5-10(-15) mm long, subjulaceous.
Pseudoparaphyllia narrow-lanceolate, entire or with
few teeth. Axillary hairs 5-6-celled, 120-140 μm long,
apical cell 45-50 × 8 μm. Stem leaves straight, 1.4-1.7
× 0.5-0.6 mm, ovate-lanceolate, concave; margins en-
tire, recurved in the upper 2/3; branch leaves 1.0-1.1
× 0.3-0.4 mm. Laminal cells 60-85 × 5-7 (4-5.5/1-
1.5) μm. Alar cells differentiated, forming a well-de-
limited excavate group with few, large, ±thin-walled,
±isodiametric cells, about 20-25 μm in diameter, which
are surrounded by 2 rows of smaller isodiametric cells.
Autoicous. Sporophytes frequent. Perichaetial
leaves to 3.0 × 0.8 mm, straight, smooth, abruptly
89
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 62. Heterophyllium affine (Hook. ex Kunth) Fleisch. (from Bolshoye Istyube Creek 470 m Ignatov 0/1181): 1
– habit; 2-4 – stem leaves; 5 – upper laminal cells of stem leaf; 6-9 – branch leaves; 10 – portion of stem with branch;
11 – capsule; 12 – stem cross-section; 13 – middle laminal cells of stem leaf; 14 – middle laminal cells of branch leaf; 15,
16 – perichaetial leaves; 17 – alar cells of stem leaf; 18 – pseudoparaphyllium; 19 – upper laminal cells of branch leaf;
20 – alar cells of branch leaf. Scale bars: 1 cm – for 1; 2 mm – for 10-11; 1 mm – for 2-4, 6-9, 15-16; 100 μm – for 5, 12-
14, 17-20.
2
15
4
13 14
1
7
10
3
5
12
8
11
17
16
18
6
19
20
9
1 cm
1 mm
2 mm
100 μm
90 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
acuminate, with indistinct costa. Seta 1.5-2.5 cm. Cap-
sule to 2.5 mm long, chestnut-brown, curved in the
upper part. Operculum high-conic to shortly oblique-
ly rostrate. Exostome teeth ca. 0.70 mm long, gradual-
ly tapered, entire; in the lower part imperfectly stri-
olate, with smooth cristae; in the middle papillose in
extensive part; in the upper part ±densely papillose;
dorsal trabeculae low, margin not incrassate; ventral
trabeculae smooth. Endostome 0.60 mm long, basal
membrane 0.20 mm high; segments rather narrow and
perforate; cilia nodose. Spores 15-17 μm.
Distribution: Callicladium haldanianum has
been reported from almost all regions of boreal and
temperate Holarctic. However, many reports from
northern and high-mountain localities are probably
based on misidentifications of Hypnum cupressi-
forme var. subjulaceum. According to our observa-
tion, C. haldanianum is a very common epixylic spe-
cies in deciduous and southern boreal forests, be-
coming much rarer and disappearing in the north-
ern taiga. It is absent also in xeric regions of Central
Asia, like Tian-Shan and Pamir Mts. and Mongolia.
In China it occurs mostly in oceanic provinces. In
Altai C. haldanianum is a rare moss restricted to
the lower elevations in vicinity of Teletzkoye Lake, a
region with especially wet and mild climate. It grows
on rotten logs, stumps and tree bases.
Specimens examined: Artybash 450 m (Zolotukhin
25.IV.1977); Bolshoye Istyube Creek 470 m (18/14a);
Chiri Creek, 0.5 km upstream 450 m (17/12; 17/13);
Klyk Pass 700 m (0/636); Kumzir 500 m (Zolotukhin
22.IV.1977); Yailyu 450 m (0/1521), 500 m (0/1522).
Heterophyllium (Schimp.) C. Müll. ex
Kindb.
The placement of Heterophyllium in Semato-
phyllaceae was argued by the deep-colored con-
cave alar group. However this character is not
impossible for Hypnaceae (cf. Hypnum cupres-
siforme var. subjulaceum, H. bambergeri, etc.).
Also, Heterophyllium was left by Tan & Buck
(1989) in Sematophyllaceae due to its relation
to Gammiella, Aptychella, Pylaisiopsis and Iso-
cladiella. However the latter four genera have
peristomes strongly different from the hypnoid
type (straight central line and heavily papil-
lose exostome, etc.), and also Aptychella has
filiform gemmae and Isocladiella a subulate
operculum. Therefore we doubt about the place-
ment of Heterophyllium near these genera in
Sematophyllaceae.
Heterophyllium includes several rather close-
ly related species, distributed in tropical to
temperate regions of both hemispheres. Plants
are autoicous and characteristically golden-yel-
low; leaves distinctly serrate above; and alar
cells dark-colored, forming an excavate group.
Heterophyllium affine (Hook. ex Kunth)
Fleisch., Musci Fl. Buitenz. 4: 1177. 1919. – Hyp-
num affine Hook. ex Kunth, Syn. Pl. Itin. Plag.
Aequin. Orbis Novi 1: 63. 1822. – Heterophyl-
lium nemorosum (Koch ex Brid.) Kindb., Ca-
nad. Rec. Sci. 6: 72. 1894. Figs. 62 & 63
Plants medium-sized to robust, yellow to brown-
ish with age. Stem 7 cm long; without hyalodermis,
with central strand; regularly pinnate-branched,
branches to 10 mm long, spreading and arcuate, the
mat thus appearing rather thick. Pseudoparaphyllia
variable, broad, triangular, narrow lanceolate to fila-
mentous. Axillary hairs 4-7-cells, ca. 150 μ m long, api-
cal cell about 45 × 8 μm. Stem leaves 2.2-2.8 × 0.6-0.7
mm, lanceolate, ±abruptly acuminate, coarsely serrate
above, somewhat plicate, gradually narrowed to the
base, orange at insertion. Branch leaves 0.8-1.4 × 0.3-
0.4 mm. Laminal cells 40-60 × 7 (4-5/2-3) μm; alar
cells subquadrate, porose and thick-walled, form-
ing a transparent to colored, sharply delimited ±ex-
cavate group.
Autoicous. Perichaetial leaves to 2.0 × 0.7 mm,
erect, smooth, serrate above; costa double or lacking.
Seta 2-3 cm, red-brown. Capsules ca. 20 mm long, in-
clined, not furrowed. Operculum conic. Annulus of 1
row of round cells, deciduous. Exostome teeth 0.5 mm
long, gradually tapered in transition zone, entire; per-
fectly striolate below; in the middle densely papillose
on many plates; in the upper part moderately papil-
lose; dorsal trabeculae and incrassate margin with
low papillae; ventral trabeculae sparsely low-papil-
lose. Endostome as high as the teeth, basal membrane
0.23 mm high, segments broad and perforate, cilia
nodose. Spores 13-15 μm.
Heterophyllium affine is easy to recognize
by the pure rich-golden color of plants, unusual
for boreal regions. Most collections in Altai were
made with abundant sporophytes.
Distribution: Heterophyllium affine occurs in tem-
perate regions of North America, northern Latin Ameri-
ca including the West Indies (in lower altitudes at higher
elevations), and in a few scattered localities in moun-
tains throughout Eurasia: Germany, Romania, Austria,
Italy, Ukrainiain Carpatians, Caucasus, Altai and Sayan
in South Siberia, Russian Far East, Bhutan in Himala-
yas, Sichuan and Hunan in mainland China, Taiwan,
and Japan. In Altai H. affine grows only around Teletz-
koye Lake, mostly in deep wet canyons, on rocks and
rotten logs, often forming pure extensive mats.
Specimens examined: Bayas Creek, in middle course 850
m (16/2); Bolshoye Istyube Creek 470 m (0/1181; 0/1183);
Kamga Creek, 10 km upstream 530 m (0/1179), 550 m (0/
1180; 0/95), 580 m (0/1182); Yailyu 450 m (0/638).
91
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1 2
3
Fig. 63. Heterophyllium affine (Hook. ex Kunth)
Fleisch. (from Kamga Creek 530 m Ignatov 0/95): 1 –
peristome (220×); 2 – outer surface of exostome in up-
per part (2200×); 3 – outer surface of exostome in lower
part (2200×).
Ptilium De Not.
The monotypic genus Ptilium is close to Hyp-
num, but differs in very dense and regularly
plumose branching, erect-ascending growth, dis-
tinctly differentiated stem and branch leaves,
linear or longly acuminate pseudoparaphyllia,
and deeply plicate, circinate leaves.
Ptilium crista-castrensis (Hedw.) De Not.,
Cronac. Briol. Ital. 2: 17. 1867. – Hypnum cris-
ta-castrensis Hedw., Sp. Musc. 287. t. 76, f. 1-4.
1801. Figs. 64 & 65
Plants large, yellow-green. Stem erect or ascend-
ing, ca. 10 cm long; without hyalodermis, with cen-
tral strand; densely plumose-branched, branches to
10 mm long; densely foliate. Pseudoparaphyllia long-
lanceolate. Axillary hairs 4-5-celled, ca. 150 μm long,
apical cell 40-50 × 8 μm. Stem leaves falcate-secund,
2.2-2.5 × 0.9-1.0 mm, lanceolate, gradually acumi-
nate, strongly curved to nearly rounded, deeply pli-
cate, serrulate above, a little narrowed at base. Branch
leaves 1.4-1.6 × 0.2-0.3 mm. Laminal cells 50-90 × 4-
6 (2-3/2-3) μm, thick-walled; alar cells few, short-
rectangular to subquadrate.
Dioicous. Sporophytes rare. Inner perichaetial
leaves to 3.2 × 0.5 mm, erect or with a little reflexed
apex, ecostate, plicate, serrulate above. Seta 2-4 cm.
Capsule 2.5 mm long, curved when open, slightly
furrowed. Operculum conic, with short truncate beak.
Annulus of one row of small round cells, deciduous.
Exostome teeth to 0.8 mm long, striolate (with pap-
illose cristae) below; in the middle densely papil-
lose; in the upper part sparsely papillose; dorsal tra-
beculae and incrassate margin with regular high-
conical papillae; ventral trabeculae sparsely tiny-
spinulose. Endostome 0.8 mm long, basal membrane
0.35 mm high; segments broad and perforate; cilia
3-4, nodose (rarely appendiculate). Spores 12-14 μm.
Distribution: Ptilium crista-castrensis is a wide-
spread circumboreal species. In Altai it occurs in
wet mossy conifer forests, and does not exceed the
tree-line in high mountains. It grows on litter among
other mosses, on rotten logs, and sometimes on rocks.
Specimens examined: Bolshoye Istyube Creek 470 m
(0/2089; 18/136); Booshkon Creek, upper course 1865 m
(Zolotukhin 2.VII.1990); Itykul Lake 1800 m (Makhatkov
& al. 2.VIII.1978); Kairu Creek, 8 km upstream 1000 m
(15/155); Kamga Creek, 11 km upstream 580 m (0/1200);
Kayakkatuyarykskij Creek 1760 m (8/76); Kishte 440 m
(Zolotukhin 11.VII.1988); Kobiguayuk Creek 2240 m (0/
605); Kuba Creek, at mouth 500 m (Voshchilko 13.VII.1970);
Ozernaya Creek 2150 m (Zolotukhin & al. 9.VIII.1976);
Seminskij Pass 1750 m (Makarov & al. 18.VI.1972); Tamanel
Peak 1800 m (34/94); Yailyu 700 m (0/606).
92 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 64. Ptilium crista-castrensis (Hedw.) De Not. (from Kayakkatuyarykskij Creek 1760 m Ignatov 8/76): 1-2 –
habit; 3 – stem cross-section; 4 – capsule; 5-7 – stem leaves; 8-10 – branch leaves; 11- middle laminal cells of stem
leaf; 12 – pseudoparaphyllia; 13 – upper laminal cells of stem leaf; 14 – alar cells of stem leaf; 15 – middle laminal
cells of branch leaf; 16 – upper laminal cells of branch leaf; 17 – alar cells of branch leaf; 18-20 – perichaetial leaves.
Scale bars: 1 cm – for 1; 5 mm – for 2, 4; 2 mm – for 18-20; 1 mm – for 5-10; 100 μm – for 3, 11-17.
17
15
20
18 19
16
13
14
1 cm
5 mm
1
5
8
6
3
4
7
910 11
12
100 μm
2
1 mm
2 mm
Gollania Broth.
Gollania was revised by Higuchi (1985),
who recognized 17 species, occurring in Hima-
layas, China, Japan, Phili ppines, Russian Far East,
and G. turgens penetrating to NW of North
America and South Siberia. Later two more
species were described from China (Higuchi &
Wu, 1995; Higuchi & Long, 1996).
The genus has an ambiguous position, since
some species have a short costa and smooth lam-
inal cells and are similar to Hypnum, while other
species with a long double costa, serrate leaf
margins, prorate cells and ±reniform leaves dem-
onstrate a closer resemblance to Rhytidiadel-
phus and Macrothamnium of Hylocomiaceae
(Higuchi, 1985). The latter family has been de-
limited by Rohrer (1985a,b) for genera which
share various combinations of the following apo-
morphic characters (1) reticulate peristome or-
namentation; (2) broad perforation of endos-
tome segments; (3) sympodial growth; (4) pres-
ence of paraphyllia; (5) differentiated stem and
93
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
1 2 3
4 5
Fig. 65. Ptilium crista-castrensis (Hedw.) De Not. (from South Siberia, Irkutsk Province, leg. L. Voronaj 1.VIII.1951,
MHA): 1 – peristome (150×); 2 – side view of exostome tooth in upper part, showing echinate-papillose inner
trabeculae (3000×); 3 – outer surface of exostome tooth in upper part (3000×); 4 – outer surface of exostome tooth in
lower part (10000×); 5 – outer surface of exostome tooth in transition zone (2300×).
branch leaves; (6) sharply serrate leave margins;
(7) costa double but long or single and branched;
(8) long laminal cells with prorate ends. Some
species of Gollania (G. ruginosa (Mitt.) Broth.,
G. varians (Mitt.) Broth., etc.) fit the characters
6, 7, 8, almost the same as Rhytidiadelphus squar-
rosus. However Gollania is different from Rhy-
tidiadelphus in: (1) plagiotropic growth; (2) ±
homomallous leaves; (3) rhizoids just below the
leaf insertion on stems and branches, while in
Rhytidiadelphus rhizoids are mostly axillary on
branch tips and on stems rhizoids are restricted
to the base of branches or branch initials; (4)
narrow perforation of endostome segments.
Gollania turgens (C. Müll.) Ando, Bot. Mag.
Tokyo 79: 769. 1966. – Cupressinia turgens C.
Müll., Nuovo Giorn. Bot. Ital., n. s., 5: 196. 1898.
Fig. 66
Plants medium-sized, yellow-green, somewhat
glossy. Stem prostrate, ca. 5 cm long; with central
strand, without hyalodermis; regularly pinnate-
branched, branches 8 mm long, ±complanate.
Pseudoparaphyllia narrowly triangular. Axillary hairs
5-celled, often all colored, 120 μm long, apical cell
35-45 × 10-12 μm. Stem leaves slightly falcate, 2.2-
2.5 × 0.6-0.8 mm, ovate-lanceolate, gradually acumi-
nate, slightly narrowed to the base, plicate; costa
double, to 1/3 the leaf length; margins serrate above,
subentire and recurved below. Branch leaves 1.4-
1.7 × 0.3-0.5 mm, widest at ca. 2/5 of leaf length.
Laminal cells (30-)40-60 × 5-7 (3-4/2-3) μm, thick-
walled, strongly prorate; alar subquadrate cells few,
gradually grading to upper cells, but abruptly de-
limited from the central basal cells.
Sporophytes not found in Altai. SEM pic-
tures of the exostome teeth have been published
by Higuchi (1985); the teeth in lower part has
papillose cristae similar to those of Ptilium.
94 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 66. Gollania turgens (C. Muell.) Ando (from Kairu Creek, 8 km upstream 1000 m Ignatov 15/2): 1-7 – habit;
2-3 – stem leaves; 4-6 – branch leaves; 8 – upper laminal cells of stem leaf; 9 – upper laminal cells of branch leaf;
10 – alar cells of stem leaf; 11 – alar cells of branch leaf; 12-13 – pseudoparaphyllia; 14 – stem cross-section; 15 –
middle laminal cells of branch leaf; 16 – middle laminal cells of stem leaf. Scale bars: 1 cm – for 7; 5 mm – for 1; 1
mm – for 2-6; 100 μm – for 8-16.
2
5
8
9
11
7
6
3
1
4
10
16
13
15
14
12
100 μm
5 mm
1 mm
1 cm
Altaian plants of Gollania turgens have the
habit of large Hypnum with slightly falcate and
plicate leaves. Higuchi (1985) mentioned that stem
leaves are typically strongly falcate, but Alaskan
plants can also have nearly straight leaves.
Distribution: A distribution map of this species was
published by Higuchi (1985). It is known from a number
of places in Alaska and northwestern Canada, but in
Asia G. turgens has very scattered localities: 2 – in
Japan, 10 – in China (Schensi, Kansu, Sichuan, Yunnan),
2 – from Nepal, 1 – from Baikal area, 1 – in Eastern
Sayan. In Altai G. turgens was collected only once, on a
wet rock outcrop near the bottom of a narrow valley.
Specimen examined: Kairu Creek, 8 km upstream 1000
m (15/2).
Taxiphyllum Fleisch.
This genus is usually placed near Isoptery-
gium s.l., Herzogiella, and Plagiothecium. Re-
95
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 67. Taxi phyllum wissgrillii (Garov.) Wijk & Marg. (from Karagai 440 m Ignatov 0/1258): 1-3 – leaves; 4-5
– perichaetial leaves; 6-7 – upper laminal cells; 8, 11 – middle laminal cells; 9,13 – alar cells; 10 – habit; 12 – stem
cross-section; 14-15 – pseudoparaphyllia. Scale bars: 5 mm – for 10; 1 mm – for 1-5; 100 μm – for 6-9, 11-15.
1 mm
100 μm
15
4
13
14
1
7
10
23
5
9
12
8
11
6
5 mm
cent raising of the weight of characters of
pseudoparaphyllia, rhizoid topography and kary-
otypes, however, demonstrates that Taxiphyllum
has little in common with the genera without
pseudoparaphyllia (Herzogiella, Isopterygiopsis
and Plagiothecium). The genus includes about 20
species in the tropical and temperate zones and is
probably not monophyletic and need a further
worldwide revision (at least exostome ornamenta-
tion was found different in some species studied).
Taxiphyllum wissgrillii (Garov.) Wijk &
Marg., Taxon 9: 191. 1960. – Hypnum wiss-
grillii Garov., Bryol. Austr. Excurs. 79. 1840.
Figs. 67 & 68
Plants small to medium-sized, green to yellow-
brown. Stem freely branched, ca. 6 cm (branches 6 mm)
when optimally developed, but usually less than 2 cm,
complanate, without hyalodermis, with central strand.
Pseudoparaphyllia narrowly triangular or lanceolate.
Axillary hairs 2-celled, 50 μm long, apical cell ca. 40 × 6
μm. Stem leaves straight, 1.0-1.5 × 0.5-0.7 mm, ovate,
±shortly acute; costa short, double or forked; margins
serrate above (teeth sometimes with recurved ti ps),
subentire below. Branch leaves 0.7-1.1 × 0.3-0.5 mm,
ovate-lanceolate. Laminal cells 60-100 × 7-9 (6-8/1)
μm, in optimally developed forms prorate on both ends
and sometimes overlapping each other because lateral
walls are not perpendicular to the lamina surface (in
poorly developed forms cells smooth). Alar cells isodi-
ametric, somewhat larger than those in the center of
the base, irregularly positioned, so the margin at base
appears somewhat eroded to toothed.
Dioicous. Sporophytes (many) found in one col-
lection from Altai (on relatively dry log on bank of
Teletzkoye Lake). Perichaetial leaves 1.0 × 0.4 mm,
plicate. Seta 0.7-1.1 cm. Capsule ca. 1.5 mm long,
inclined, slightly curved. Exostome teeth ca. 0.50-
0.53 mm long, ±gradually tapered, slightly serrulate
to subentire above; in the lower part striolate, with
smooth cristae; in the middle with short irregular
vermicular cristae (often longitudinally oblique)
sometimes intermixed with papillae; in the upper
96 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
7
1 2 3
4 5
6
Fig. 68. Taxi phyllum wissgrillii (Garov.) Wijk & Marg. (from Karagai 440 m Ignatov 0/1258): 1 – peristome
(150×); 2 – side view of exostome tooth in upper part (3900×); 3 – outer surface of exostome tooth in upper part
(4400×); 4-5 – upter surface of exostome tooth in transition zone (4 – 2750×; 5 – 3600×); 6 – outer surface of
exostome tooth just below transition zone (5600×); 7 – outer surface of exostome tooth in lower part (3300×).
part abruptly becoming smooth, with rare low papil-
lae only on strongly incrassate dorsal trabeculae;
margin non-incrassate; ventral surface and trabecu-
lae smooth. Endostome 0.55 mm long, basal mem-
brane 0.2 mm high; segments broad and perforate;
cilia nodose. Spores. 12-16 μm.
Most specimens of this species from north-
ern Europe are sterile and represent a rather
small procumbent form. Such plants occur also
in Altai, especially in shady rock crevices, where
97
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
scattered shoots can be collected. However, sev-
eral collections differ in larger size, subjula-
ceous foliage, growth on rather dry rotten logs
in diffuse shade, and sometimes in having sporo-
phytes. These plants fit well the type of Isop-
terygium depressum var. subjulaceum Card. et
Coppey, Bull. Soc. Bot. France 58: 198. 1911. (?iso-
type: pierres calcaires pres Nancy, France, 4 Mars
1911, leg. Coppey, H-BR!). Identical plants were
seen from South Europe (Hungary, Italy) and
Kazakhstan. However their taxonomic recogni-
tion seems unreasonable since (1) leaf shape and
areolation of larger and smaller form are simi-
lar in essential features; (2) transitional forms
have been observed in South Europe and Cau-
casus (in H).
Distribution: This species has a mostly Europe-
an distribution, known from nearly all countries and
reaching to the east Ural Mts. and Caucasus. Out-
side Europe it has been reported from North Africa,
Himalayas and Altai Mts. In Altai T. wissgrillii
grows mostly on deeply shaded rocks (usually ±cal-
careous schists), rarely occurring on dry logs among
thick moss mats, in partial shade.
Specimens examined: Artybash 450 m (22/4); Ayukol
800 m (0/1253); Chemal Creek, 3 km upstream 450 m
(34/106; 34/27); Derbogach 440 m (0/13); Elekmonar
Creek, 8 km upstream 800 m (26/66); Karagai 440 m
(0/1258; 0/276); Kobukhta 550 m (0/607), 600 m (0/
1255); Kumzir 450 m (20/13; 20/27); Maima 350 m
(35/47); Ok-Porok Creek, at mouth 450 m (0/1259; 0/
1260); Tura Creek, in middle course 1300 m (28/35);
Ust-Sema 450 m (24/86); Yailyu 450 m (0/1263), 550
m (0/1257; 0/1261), 600 m (0/1256); Yurga 540 m
(0/1254)
Sematophyllaceae Broth.
The family is a large and heterogeneous as-
semblage of tropical to temperate ecostate pleu-
rocarps characterized by (1) angular cells of-
ten vesiculate-inflated; (2) tendency to peris-
tome reduction, including both exostome (teeth
from cross-striolate to heavily papillose, with
straight, often furrowed center line) and en-
dostome (cilia, basal membrane and sometimes
segments lacking). The family includes about
40 genera, and not in all of them are both ga-
metophytic and sporophytic characters corre-
lated with each other. Not rare the character
either 1, or 2 are considered enough to refer a
moss to Sematophyllaceae.
Struckia C. Müll.
This genus includes one species with two
subspecies. It is peculiar in lax areolation in
leaf base, fragile branches, single peristome with
a straight median line (with thickenings along
the median line on the ventral surface of peris-
tome). Tan & Buck (1989) transfer Struckia
from Sematophyllaceae (where it was placed
by Brotherus, 1925) to Hypnaceae because of
unusually lax areolation of leaves. Lax leaf ar-
eolation, fragile branches and rhizoids arising
on leaf back could be interpreted as characters
of Plagiotheciaceae, where Struckia was placed
by Fleischer (1912). However lack of endos-
tome and relatively short, blunt and strongly
papillose exostome teeth are not known in any
genera of both Hypnaceae and Plagiotheciace-
ae, while the similar pattern of peristome reduc-
tion is known in many genera of Sematophyl-
laceae. In some species of Sematophyllaceae, for
example Brotherella recurvans from North
America, we found rather fragile branches and
rhizoid position similar to Struckia (mostly
on the back of leaf at base). Sematophyllaceae
are badly waiting for revision, and probably
Struckia is not close to the main nucleus of this
family, but the placement of this genus in Hyp-
naceae seems equally inappropriate. We leave
here this problem for further studies. Moreover
Siberian plants have never been found with
sporophytes.
Struckia argentata (Mitt.) C. Müll. ssp. ze-
rovii (Lazarenko) Tan, Buck et Ignatov, Lind-
bergia 16(3): 102. 1990. – Cephalocladium ze-
rovii Lazarenko, Bot. Zhurn. AN URSR 3(3-4):
62. 1946. Fig. 69
Plants rather small, in soft green to whitish-green
±glossy mats. Stem prostrate, to 1 cm long, freely
branched, julaceous, with some stoloniform rarely
foliated axes. Many shoots ending in crown of small
caducous leaves 0.3-0.5 × 0.2 mm and naked axes 1-
2 mm below them. Pseudoparaphyllia lanceolate.
Axillary hairs 2-3-celled, ca. 170 μm long, apical cell
65 × 7.5 μm. Leaves straight, 0.9-1.5 × 0.2-0.4 mm,
lanceolate, acuminate, entire to indistinctly serrulate
above. Laminal cells 60-110(-145) × 10-16(-20 μm).
Alar cells wider and shorter, in wide group across
the leaf base. Dioicous. Sporophytes unknown in this
subspecies.
Lazarenko (1946) described Cephalocladi-
um zerovii from Altai as a new species and a
new monotypic genus. Bardunov (1974) found
it ± widespread in South Siberia, eastward to
Baikal Lake. Abramova & Abramov (1981)
found this species in Mongolia and demon-
98 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 69. 1-10, 12-14, 16 – Struckia argentata ssp. zerovii (Lazarenko) Tan, Buck et Ignatov (from Yailyu 450 m
Ignatov 0/1185) and 11, 15 S. argentata C. Müll. ssp. argentata (from Buthan, Griffith LE): 1-3 – stem leaves; 4,14
– caducous leaves; 5 – stem cross-section; 6-9 – habit; 10, 11 – middle laminal cells of stem leaf; 12 – upper laminal
cells of stem leaf; 13 – alar cells of stem leaf; 15, 16 – leaf cross-section. Scale bars: 2 mm – for 6-9; 0.5 mm – for 1-
4; 100 μm – for 5, 10-16.
6
10
15
1
23
4
5
7
89
11
12
13
14
100 μm
0.5 mm 2 mm
16
strated that Siberian and Mongolian plants
are the same as Fabronia enerve Broth. de-
scribed from Sichuan. Tan & al. (1990) com-
pared Cephalocladium zerovii and Fabronia
enerve with Struckia argentata, a monotypic
genus known from Himalayas and Yunnan, and
found them conspecific. However they found
that southern population of S. argentata dif-
fers from the northern by (1) tumid appear-
ance; (2) distinctly serrate leaves; (3) longer
cells; (4) frequent sporophytes production;
and (5) absence of caducous leaves. These dif-
ferences were evaluated as subspecific because
in the easternmost locality of the southern
population in Sichuan intermediate plants was
found (Fabronia enerve).
99
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Subsequent new collections from Sichuan
(Koponen 46069, 45584, 46469 H!) also repre-
sent plants with a fairly long naked subtermi-
nal zone and nearly entire leaves, exactly iden-
tical with the Altaian. Also more collections from
Altai revealed that the cell length is too vari-
able to be an useful character. Also, one pecu-
liarity of the Himalayan plants was neglected
– laminal cell ends often overlap each other,
resulting in unclear cell outlines in front view,
and specifically a roughened lamina surface seen
in cross section (Fig. 000). We leave here the
taxonomy of Struckia unchanged, since it will
be discussed in a separate paper.
Distribution: Struckia argentata ssp. zerovii is
rather widespread at lower elevation on mountains
of South Siberia, from Altai to Baikal area, Northern
Mongolia and Sichuan. In Altai it grows usually on
rock outcrops, rarely on trunks, in the forest zone,
but once collected in the subalpine zone.
Specimens examined: Chemal Creek, 3 km upstream
450 m (34/49); Chiri Creek, 0.5 km upstream 450 m
(17/9); Chulcha River, in middle course 960 m (9/142),
1000 m (9/29), 1100 m (9/41; 9/50); Elekmonar Creek,
5 km upstream 700 m (26/27); Kairu Creek, 8 km up-
stream 1000 m (15/66); Oligash (between Katun and
Sema) m (Zerov 16.VIII.1940 LB!); Yailyu 440 m (0/
1190; 0/637), 445 m (0/1186), 450 m (0/1185; 0/1187;
1/13; 1/5; 1/6), 460 m (0/1188; 0/1189), 580 m (0/
1184); Yakhansoru Lake 1850 m (Zolotukhin 28.VI.1990);
Yurga 450 m (21/43).
Hylocomiaceae (Broth.) Fleisch.
The contents of this family was discussed
by Rohrer (1985a,b). He concluded that though
there is no one character to delimit Hylocomi-
aceae from Hypnaceae, the former probably com-
prises a natural segregate. The apomorphic char-
acters of Hylocomiaceae shared by only some
of its representatives are as follows: (1) many
species robust, forming a carpet in boreal for-
ests or subalpine communities; (2) in Pleuro-
zium and Rhytidiadelphus stems often ascend-
ing to erect in dense growth; in loose growth
forms numerous sympodial branches are devel-
oped, which are always numerous in Hylocomi-
um and Hylocomiastrum; (3) in all four gen-
era rhizoids mostly on branch tips, rare or lack-
ing on the stem; (4) Hylocomium and Hyloco-
miastrum have paraphyllia; (5) in Hylocomi-
astrum, Hylocomium and Rhytidiadelphus
leaves are coarsely serrate; (6) in Hylocomias-
trum umbratum and Rhytidiadelphus triquetrus
costa is long and double; (7) in Hylocomium
and Rhytidiadelphus triquetrus laminal cells
are often prorate; (8) Pleurozium schreberi, Hy-
locomium splendens and Rhytidiadelphus tri-
quetrus have reticulate ornamentation in the
lower exostome teeth; (9) Pleurozium, Hyloco-
mium and Rhytidiadelphus have endostome
segments broadly perforate.
Hedenäs (1989) mentioned for Hylocomi-
aceae rhizoids on the stem just below the leaf
insertion and on branch tips in axillary posi-
tion. We have seen rhizoids on the stem below
leaves only in both Altaian species of Hyloco-
miastrum. In Rhytidiadelphus squarrosus they
were found only around branch initials. In R.
triquetrum, Pleurozium and Hylocomium no
rhizoids on the stem were observed. On branch
tips rhizoids are ±clearly axillary or in ±indef-
inite position in between leaves.
Hylocomiastrum Fleisch.
This genus is sometimes not segragated from
Hylocomium, since both genera have abundant
paraphyllia and robust plants forming exten-
sive mats. Rohrer (1985b) summarized the dif-
ferences between Hylocomiastrum and Hylo-
comium as follows: (1) branching pattern vari-
able vs. regularly bipinnate and sympodial; (2)
paraphyllia of rain-deer type vs. deer-horn type;
(3) leaves strongly plicate; (4) costa stronger;
(5) operculum conic vs. rostrate; (6) exostome
striolate below vs. reticulate; (7) segments nar-
rowly perforate.
Hylocomiastrum includes 3 species, 2 of
which has wide circum-holarctic distribution,
and H. himalayanum (Mitt.) Broth. is a moun-
tainous East Asian species.
KEY TO THE ALTAIAN SPECIES OF
HYLOCOMIASTRUM:
1. Plants 1(-2)-pinnate, glossy, pale; leaves non-
decurrent; costa single . . . H. pyrenaicum
1. Plants (1-)2-3-pinnate, dull, dark-green;
leaves decurrent; costa double
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. umbratum
Hylocomiastrum pyrenaicum (Spruce)
Fleisch. in Broth., Nat. Pflanzenfam., ed. 2, 11: 487.
1925. – Hypnum pyrenaicum Spruce, Musci Pyren.
4. 1847. – Hylocomium pyrenaicum (Spruce)
Lindb., Musci Scand.: 37. 1879. Fig. 70
Plants robust, in pale-green, somewhat glossy
mats. Stem arcuate, regularly pinnate, 7 cm long;
epidermal cells thick-walled, central strand weak.
100 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Fig. 70. Hylocomiastrum pyrenaicum (Spruce) Fleisch. (from Karakol Lakes 1900 m Ignatov 28/87): 1 – stem
portion; 2-4 – stem leaves; 5-7 – branch leaves; 8 – habit; 9 – middle laminal cells of stem leaf; 10 – upper lamina
cells of stem leaf; 11 – alar cells of stem leaf; 12 – proximal branch leaf; 13-14 – paraphyllia; 15 – stem cross-section.
Scale bars: 1 cm - for 8; 2 mm – for 1; 1 mm – for 2-7; 100 μm – for 9-15.
4
13
1
7
10
35
9
8
11
26
100 μm
15
14
12
2 mm
1 mm
1 cm
Paraphyllia abundant on stem and branches, dev-
ided mostly to 2-3 branches from the base, but para-
phyllia around branch primordia longer, irregular-
ly pinnate to bipinnate branching. Pseudoparaphyl-
lia transversely ovate at base of branch primordia.
Axillary hairs 4-5-celled, 100-150 μm long; apical
cell 50 × 11 μm. Stem leaves erect, 1.8-2.2 × 0.8-1.0
mm, broadly ovate, acuminate, rounded at base, non-
decurrent, plicate; costa single, up to 3/4 the leaf
length; margins recurved at base, coarsely serrate
above, subentire below. Laminal cells 30-55 × 5-6.5
(4-4.5/1.5-2) μm. Cells in leaf corners not differ-
entiated.
Dioicous. No sporophytes found in Russian Al-
tai, but old setae were seen in one collection from
Kazakh Altai (Leninogorsk Distr., 1300 m, Voro-
schilov 3608, MHA). Perichaetial leaves 1.6 × 0.8
mm, clasping the base of seta, with reflexed apex,
ecostate, smooth.
Distribution: The species is common in moun-
tainous areas throughout northern parts of the Hol-
arctic, growing in boreal and alpine types of vegeta-
101
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 71. Hylocomiastrum umbratum (Hedw.) Fleisch. (from Uedinennoye Lake 880 m Ignatov 0/604): 1 – habit;
2 – capsule; 3 – upper laminal cells of stem leaf; 4-6 – stem leaves; 7-8 – branch leaves; 9-10 – outer perichaetial
leaves; 11 – inner perichaetial leaf; 12 – portion of stem with branches; 13-14 – pseudoparaphyllia; 15 – alar cells
of stem leaf; 16 – stem cross-section; 17-18 – paraphyllia; 19 – paraphyllium of close proximity to branch primordia;
20 – middle laminal cells of stem leaf. Scale bars: 1 cm – for 1; 2 mm – for 2, 12; 1 mm – for 4-9; 0.5 mm – for 10-
11; 100 μm – for 3, 13-20.
12
9
2
15
4
13
14
1
7
10
3
5
8
11
17 18
6
19
16
100 μm
20
1 cm
2 mm
1 mm
2 mm
tion. In Altai Hylocomiastrum pyrenaicum occurs
mostly in the alpine and subalpine zones, on soil under
shrubby and herbaceous canopy and among rocks.
In the forest zone it grows sporadically on rock out-
crops or on organic substrates (litter, old stumps,
tree bases).
Specimens examined: Ak-Turu Creek, upper course 2500
m (Nekrasova 11.VIII.1911 LE!); Ayukol 1350 m (0/
1212; 0/596); Bayas Lake 1750 m (0/1217), 2000 m (0/
1213; 0/597); Bolshoye Kurkure Creek, upper course 2300
m (Galanin 9.VIII.1976); Kairu-Bazhi Peak 2150 m (13/
18); Kaitanak Creek 1500 m (A. Krylov & Rechan
10.VII.1968 LE!); Karakol Lakes 1900 m (28/132; 28/
102 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 2
4 53
Fig. 72. Hylocomiastrum umbratum (Hedw.) Fleisch. (from Uedinennoye Lake 880 m Ignatov 0/604): 1-2 –
peristome (1 – 150×; 2 – 225×); 3 – outer surface of exostome tooth in upper part (2250×); 4 – side view of exostome
tooth in upper part, showing incrassate and warty papillose inner trabeculae (4000×); 5 – outer surface of exostome
tooth in lower part (4000×).
162; 28/62; 28/87); Kayakkatuyarykskij Creek 2150 m
(7/18); Kobiguayuk Creek 2400 m (0/599; 0/600);
Sarulu Creek, upper course (Nikitina 30.VI.1925 LE!);
Srednij Shaltan Creek 600 m (0/1216), 620 m (0/1218),
670 m (0/1214); Tabozhok Peak 2600 m (31/41); Tosh-
kalykaya Peak 2500 m (Zolotukhin 14.VII.1990);
Uedinennoye Lake 880 m (0/1215).
Hylocomiastrum umbratum (Hedw.)
Fleisch. in Broth., Nat. Pflanzenfam., ed. 2, 11:
487. 1925. – Hypnum umbratum Hedw., Sp.
Musc. Frond. 263. t. 67 f. 10-13. 1801. – Hyloco-
mium umbratum (Hedw.) B. S. G., Bryol. Eur.
5: 175. 1852. Figs. 71 & 72
Plants robust, in dull dark-green loose mats. Stem
to 10 cm long, once, bi- or tripinnate-branched , ±ar-
cuate, with thick-walled epidermal cells and weak
central strand. Paraphyllia densely covering the stem
and branches, strongly branched from the base, but
paraphyllia around branch primordia longer, pin-
nate-branching to serrate-ciliate. Pseudoparaphyllia
transversely ovate. Axillary hairs 3-4-celled, ca. 150
μm long, apical cell ca. 60 × 10 μm. Stem leaves loose-
ly erect, 1.5-1.8 × 1.0 mm, broadly ovate-triangular,
shortly acuminate, narrowly decurrent; costa dou-
ble, reaching 1/3–2/3 of the leaf length; margins
coarsely serrate. Branch leaves smaller, ovate-lan-
ceolate. Laminal cells 30-60 × 4.5-7 (3-5/1.5-3) μm,
±smooth. Cells in basal corners not differentiated.
Dioicous. Sporophytes rather rare. Outer pericha-
etial leaves 3.2 × 1.0 mm, smooth, with costa double
103
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 73. Hylocomium splendens (Hedw.) B.S.G. (from Bolshoye Istyube Creek 500 m Zolotukhin 19.V.1977): 1-3 –
stem leaves; 4 – upper lamina cells of stem leaf; 5 – capsule; 6-9 – branch leaves; 10 – middle lamina cells of stem
leaf; 11- habit; -12 – alar cells of stem leaf; 13-14 perichaetial leaves; 15 – stem portion; 16 – stem cross-section; 17-
18 – proximal branch leaves; 19-20 – paraphyllia; 21 – stem portion with branches. Scale bars: 1 cm – for 11; 5 mm
– for 5, 21; 2 mm – for 15; 1 mm – for 13-14; 0.5 mm – for 1-3, 6-9; 100 μm – for 4, 10, 12, 16-20.
13
15
4
13 14
7
10
5
8
11
12
9
17
2
18
6
19
16
21
20
1 cm
5 mm
2 mm
0.5 mm
1 mm
100 μm
or single, sometimes forked, to 2/3 the leaf length,
serrate above; inner perichaetial leaves shorter, re-
flexed, with weak double costa. Seta 3 cm. Capsule ca.
2.5 mm long. Operculum conic. Exostome teeth 0.60
mm long, abruptly tapered in the transition zone,
subentire above; perfectly reticulate below; in the
transition zone densely papillose (few plates); in the
upper part moderately low-papillose; dorsal trabec-
ulae rather high; margin not especially incrassate,
slightly papillose; ventral trabeculae strongly incras-
sate and papillose. Endostome as long as the teeth,
basal membrane ca. 0.3 mm high; segments widely
perforate; cilia nodose, sometimes appendiculate.
Spores 14 μm.
Distribution: Hylocomiastrum umbratum has a
scattered distribution in oceanic and mountain ar-
eas of the Holarctic, with few scattered localities in
southern parts of the boreal zone on lowlands. In
Altai it was found few times in deep canyons or in
wet fir forests, at lower elevations.
Specimens examined: Bolshoye Istyube Creek 470 m
(18/108); Kamga Creek, upper course 850 m (Martynen-
ko 9.VIII.1935 LE!); Uedinennoye Lake 880 m (0/1210;
0/604), 1030 m (0/1211).
104 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 2
3
Fig. 74. Hylocomium splendens (Hedw.) B.S.G. (from
Bolshoye Istyube Creek 500 m Zolotukhin 19.V.1977):
1 – peristome (200×); 2 – outer surface of exostome
tooth in upper part (3000×); 3 – outer surface of exos-
tome tooth in lower part (1100×).
Hylocomium B. S. G.
The genus includes one species widely dis-
tributed in cool climates.
Hylocomium splendens (Hedw.) B.S.G.,
Bryol. Eur. 5: 173. 1852. – Hypnum splendens Hedw.,
Sp. Musc. 262. t. 67., f. 7-9. 1801. Figs. 73 & 74
Plants robust, pale-green to brownish-green. Stem
to 15 cm long, with weak central strand, monopodially
bipinnate-branched and also regularly sympodially
branched, resulting in step-wise appearance. Paraphyl-
lia densely covering stems and branches. Pseudopara-
phyllia broadly to transversely ovate. Axillary hairs 3-
5-celled; ca. 100 μm long, apical cell 40 × 10 μm. Stem
(i. e., main sympodial branch) leaves tightly clasping
stem at base, ±flexuose above, ca. 2.0-2.5 × 1 mm, ovate-
triangular, abruptly narrowed to flexuose-rugose acu-
men, slightly plicate; costa double or single, to 1/3(-
1/2) the leaf length, margins serrate. Branch leaves
smaller, oblong-lanceolate. Laminal cells 30-60 × 5 (3/
2) μm, thick-walled, prorate. Cells in leaf corners not
differentiated, across the base orange-colored.
Dioicous. Sporophytes rather rare. Perichaetial leaves
to 3.5 × 1.0, erect, ecostate. Seta 2-3 cm. Capsule 2.0-2.5
mm long. Operculum obliquely rostrate. Exostome teeth
0.6 mm long; abruptly tapered in the transition zone,
subentire above; in the lower part imperfectly reticu-
late (transverse cristae wavy and anastomozed); in the
transition zone densely papillose on few plates; in ther
upper part moderately low papillose, with rare low
papillae on relatively low dorsal trabeculae; margin
incrassate, slightly papillose; ventral trabeculae smooth.
Endostome about 0.55 mm long, basal membrane 0.2
mm high; segments broad and widely perforate; cilia
nodose. Spores about 15 μm.
In Altai Hylocomium splendens is not espe-
cially polymorphous, except one form found in
abundance in shrubby hummocks in a flood
valley in the upper taiga zone, where plants have
erect growth, less developed secondary branch-
es and totally lacking sympodial branching.
These characters fit the common Arctic var. ob-
tusifolium (Geh.) Par. (= H. alaskanum Lesq.
et James) Aust. However it is not apparent
whether this population was derived from non-
Altaian source or originated by means of a par-
allel mutation in Altaian plants.
Distribution: Very common in the Arctic
and boreal regions of the Holarctic and also in
conifer forests in more southern mountains, in-
cluding Central and South America, Africa and
New Zealand. In Altai it is very common, being
absent only in very xeric regions.
105
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
100 μm
Fig. 75. Rhytidiadelphus squarrosus var. calvescens (Lindb.) Warnst. (from Yailyu 450 m Ignatov 0/1219): 1-2 –
stem leaves; 3-5 – branch leaves; 6 – middle laminal cells of stem leaf; 7 – alar cells of stem leaf; 8 – habit; 9 –
upper laminal cells of stem leaf; 10-11 – pseudoparaphyllia; 12 – stem cross-section. Scale bars: 1 cm – for 8; 2 mm
– for 1-5; 0.2 mm – for 10-11; 100 μm – for 6-7, 9, 12.
1 cm
1
5
8
6
34
7
910 11 12
0.2 mm
2
2 mm
Specimens examined: Ayulyuyuzyuk Creek 2100 m (0/
173); Bayas Lake 1750 m (0/603); Bele 500 m (Zolo-
tukhin 28.VII.1982); Bogoyash Creek, at mouth 2070 m
(Zolotukhin & al. 17.VIII.1977); Bolshoye Istyube Creek
500 m (Zolotukhin 19.V.1977); Chibitka Creek, at mouth
2200 m (Kuklina VII.1984); Kamga Bay 450 m (Zolo-
tukhin 20.X.1988); Karatash 450 m (Zolotukhin & al.
26.V.1984); Kishte 440 m (Zolotukhin 11.VII.1988);
Kobiguayuk Creek 2908 m (0/601); Maloye Istyube Creek,
1 km upstream 600 m (Zolotukhin 19.V.1977); Ozernaya
Creek 2150 m (Lebedeva & al. 9.VIII.1978); Saratki Cape
440 m (Zolotukhin 14.X.1987); Seminskij Pass 1750 m
(Makarov & al. 18.VI.1972); Tokpak Creek, in middle
course 2600 m (36/387a); Yailyu 450 m (0/602).
Rhytidiadelphus (Lindb. ex Limpr.) Warnst.
The genus includes 4-5 species, distributed
in boreal and hemiboreal regions of the Hol-
arctic. Main characters of the genus include:
plants robust, without paraphyllia; leaves re-
flexed to squarrose, ±ovate, acuminate, serru-
late to serrate.
Rhytidiadelphus triquetrus differs from the
rest of the species in the genus by (1) rigid
straight leaves; (2) long costa; (3) prorate cells;
(4) reticulate exostome; and (5) unusually nar-
row axillary hairs. These seem enough for a new
genus, but the conservative position seems more
preferable, since the closest relatives of R. tri-
quetrus are other members of the same genus.
KEY TO THE ALTAIAN SPECIES OF
RHYTIDIADELPHUS
1. Leaves gradually narrowed upward; double
costa reaching about 3/4 the leaf length;
laminal cells strongly prorate . . R. triquetrus
1. Leaves ±abruptly narrowed upward; dou-
ble costa reaching 1/3-1/2 the leaf length;
laminal cells smooth
. . . . . . . . . . . . . . . R. squarrosus var. calvescens
Rhytidiadelphus squarrosus (Hedw.)
Warnst., Warnst., Krypt.-fl. Mark Brandenb. 2:
918. 1906. var. calvescens (Lindb.) Warnst.,
Krypt.-fl. Mark Brandenb. 2: 920. 1906. – Rhy-
tidiadelphus subpinnatus (Lindb.) T. Kop.,
Hikobia 6: 19. 1971. – Hylocomium subpinna-
tum Lindb., Hedwigia 6: 41. 1867. Fig. 75
106 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
Plants robust, in loose green mats. Stem ascend-
ing to somewhat arcuate above, to 12 cm long; with
weak central strand; ±regularly pinnate-, sometimes
bipinnate-branched. Pseudoparaphyllia broadly tri-
angular. Axillary hairs 4-6-celled, 150-200 μm long,
apical cell 50 × 8 μm. Stem leaves squarrose, 2.2-2.5
× 1.3-1.5 mm, ovate base appressed to erect, upper
part reflexed; costa double, up to 1/5-1/4 the leaf
length; margins serrate above, serrulate below. Prox-
imal branch leaves 2.0 × 0.9-1.0 mm. Median lami-
nal cells 50-80 × 7-8 (6-7/1-1.5) μm, smooth, near
the base at margin ca. 40 × 7-8 μm, between leaf cor-
ners and costa cells larger, ca. 40 × 20 μm.
Dioicous. No sporophytes found in Russian Al-
tai. SEM pictures of the striolate exostome ornamen-
tation and of the endostome have been published by
Rohrer (1985b).
The differences between typical Rhytidia-
delphus squarrosus and R. squarrosus var.
calvescens (= R. subpinnatus) have been eval-
uated differently. American bryologists have
consided R. squarrosus in a wide sense (Crum
& Anderson, 1981; Rohrer, 1985), while most
European authors consider them as separate
species. Important for their distinction are:
var. squarrosus: plants in dense extensive
mats; stem tips straight and stellate due to
strongly squarrose leaves; branches irregular
and relatively short; stem leaves reflexed to
squarrose, ovate at base and clasping the stem;
lower branch leaves of the same shape, finely
serrulate.
var. calvescens: plants in loose mats, often
distally arcuate; ±regularly pinnate-branched,
branches sometimes with secondary branches;
stem leaves reflexed, but not squarrose, broad-
est near the base, so leaves are not clasping the
stem which is well visible in between leaves;
lower branch leaves different from stem leaves,
rather coarsely serrate.
All the specimens from Altai belong to the
latter variety.
Though typical expressions of these variet-
ies look very distinct, transitional forms occur
in herbaria. Field observations in Ural Mts. and
European Russia showed that in woodlands on
bogging clear-cut areas and along wet trails
var. calvescens (common in nearby forests) is
gradually modified towards var. squarrosus,
becoming sometimes indistinguishable from
North-European plants. High moisture and open
light are probably responsible for this change.
Some Altaian plants from open places in subal-
pine zone also are somewhat similar to R. squar-
rosus s. str. in habit.
It is highly probable that the var. squarrosus s.
str., a weed of grasslands in North Europe, com-
prises a monophyletic line. Rohrer (1985) consid-
ered it to have originated in disturbed grass-
lands, while Koponen (1971) suggested that its
original habitats were sea shore meadows. How-
ever there is virtually no way to distingush it
from “squarrosus-like” expressions of var. calve-
scens in Chukotka, Alaska, British Columbia, moun-
tains of South Siberia and East European low-
lands. The only difference is that North-European
var. squarrosus forms extensive populations, which
are very constant in their characters, while in large
populations from other places trends to var. calve-
scens are usually apparent.
Distribution: Rhytidiadelphus squarrosus var.
calvescens has a Holarctic distribution confined
mostly to boreal forests, penetrating in coniferous
forest in mountains of Central Europe, Hokkaido
and Honsu in Japan, northern parts of mountains of
Middle Asia. Reported from Sichuan. In Altai it has
scattered distribution in conifer forests, under tall
herbs (both in the forest and subalpine belts), and
in dwarf Betula shrubs in the subalpine zone.
Specimens examined: Bardaky Lake 2050 m (13/112);
Bayas Creek, in middle course 1100 m (16/16); Bolshoye
Istyube Creek 470 m (18/131); Gorno-Altaisk 400 m (35/
53); Kairu Creek, 22 km upstream 1550 m (14/31);
Kaitanak Creek 1500 m (A. Krylov & Rechan 10.VII.1968
LE!); Oligash (between Katun and Sema) (Zerov
17.VIII.1940 LB!); Uedinennoye Lake 880 m (0/1220);
Ust-Sema 500 m (24/14); Yailyu 450 m (0/1219), 600
m (0/1221).
Rhytidiadelphus triquetrus (Hedw.)
Warnst., Krypt.-Fl. Brandenburg, Laubm. 920: 996.
t. 926 f. 6. 1906. – Hypnum triquetrum Hedw.,
Sp. Musc. Frond. 256. 1801. Figs. 76 & 77
Plants robust, in yellow-green mats. Stem ascend-
ing, to 10 cm long; with weak central strand; regu-
larly pinnate-branched, leaves and young branches
crowded terminally, so stem is subcomose, branches
1.7 cm long, attenuate at tips. Pseudoparaphyllia
broadly triangular. Axillary hairs 4-6-celled, ca. 200
μm long, apical cell 45-50 × 6 μm. Stem leaves ±re-
flexed to wide spreading, 4.0 × 2.1 mm, ovate-trian-
gular, cordate at base, gradually acute, plicate; costa
double, long, 3/4-4/5 the leaf length; margins ser-
rate. Branch leaves 2.3-2.4 × 0.9-1.0 mm, of the same
shape as stem leaves. Laminal cells 60-110 × 7 (3-4/
2-3) μm, strongly prorate. Alar cells subquadrate, form-
ing small indistinct group.
107
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
Fig. 76. Rhytidiadelphus triquetrus (Hedw.) Warnst. (from Kairu Creek, 8 km upstream 1000 m Ignatov 15/49):
1 – stem leaf; 2 – upper lamina cells of stem leaf; 3-4 – branch leaves; 5 – middle lamina cells of stem leaf; 6 – habit;
7 – alar cells of stem leaf; 8-9 – perichaetial leaves; 10-11 – pseudoparaphyllia. Scale bars: 1 cm – for 6; 2 mm – for
1, 3-4, 8-9; 0.2 mm – for 10-11; 100 μm – for 2, 5, 7.
4
1
710
2
3
5
9
8
2 mm
0.2 mm
100 μm
11
1 cm
6
Dioicous. Sporophytes rather rare. Perichaetial
leaves 3.3-4.1 × 0.7-1.2 mm, oblong, abruptly acumi-
nate, smooth, entire, with indistinct double costa to
about 1/4 the leaf length. Seta 1.5-2.0 cm. Capsule
2.0-2.5 mm long, furrowed when open and dry. Exos-
tome teeth 0.6 mm high; straight to ca. 2/3, distinctly
narrowed in the transition zone, entire, regularly re-
ticulate below; in the middle abruptly grading to the
upper part where smooth or longitudinally furrowed;
dorsal trabeculae prominent, with incrassate margin;
ventral trabeculae smooth. Endostome 0.55 mm long,
basal membrane ca. 0.30 mm high; segments broadly
perforate and usually strongly split; cilia nodose, some-
times eroded-flexuose. Spores 18-23 μm.
Distribution: The species is common in the bore-
al and hemiboreal zones and at less extend in tem-
perate regions of the Holarctic, and in Himalayas. In
Altai it is common in ±mesic forests, but never pen-
etrates above tree-line.
Specimens examined: Adylda Creek 1100 m (34/140;
34/82); Ayulyuyuzyuk Creek 2100 m (0/383); Bardaky
Lake 2050 m (13/113); Bolshoi Yaloman Creek, in mid-
dle course 1500 m (Makarov & al. 15.VII.1972); Bolshoye
Istyube Creek 470 m (0/2089a; 18/132); Chelyush Cape
450 m (Zolotukhin & al. 19.VII.1980); Chemal Creek, 3
km upstream 450 m (34/25); Chemal, 5 km to the SE 500
m (Makarov & al. 27.VI.1972); Kairu Creek, 8 km up-
stream 1000 m (15/135a; 15/49); Kairu-Bazhi Peak 2050
m (13/107a); Karagai 450 m (Zolotukhin 11.VIII.1988);
Kayakkatuyarykskij Creek 1600 m (8/69); Kishte 440 m
(Zolotukhin 11.VII.1988); Korbu 440 m (Zolotukhin
20.X.1988); Malyi Yaloman Creek, 8 km upstream 900 m
(25/22); Yailyu 450 m (0/1204), 500 m (0/1203).
Pleurozium Mitt.
This monotypic genus has a long history of
placement in various families (see a review in
Rohrer, 1985a,b). The placement in Entodonta-
ceae was suggested by Fleischer (1914, 1923)
and Brotherus (1925), and was accepted in many
manuals. Some authors placed it in Hypnaceae
without certain relations to other groups (Nish-
imura & al., 1984). Another position, close to (or
within) Hylocomium and Rhytidiadelphus, was
accepted by De Notaris (1867), Limpricht (1895-
1904), Nyholm (1965) and in recent literature
also by Buck (1980) and Rohrer (1985a, b).
Important characters includes robust plants, form-
ing extensive mats in conifer forests, rhizoids
mostly on branch tips, reticulate exostome orna-
mentation, and broad perforation of endostome.
108 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 2
3 4
Fig. 77. Rhytidiadelphus triquetrus (Hedw.) Warnst. (from Kairu Creek 1000 m Ignatov 15/49): 1 – peristome
(160×); 2 – side view of exostome tooth in upper part, showing smooth inner trabeculae (3500×); 3 – outer surface of
exostome tooth in lower part (5500×); 4 – outer surface of exostome tooth in transition zone (4200×).
Plerozium differs from related genera by (1)
lacking paraphyllia; (2) leaves obtuse and nearly
entire; (3) ±distinctly differentiated alar groups.
Pleurozium schreberi (Brid.) Mitt., J. Linn.
Soc., Bot. 12: 537. 1869. – Hypnum schreberi Brid.,
Muscol. Recent. 2(2): 88. 1801. Figs.78 & 79
Plants rather robust, in green to yellow-green
glossy extensive mats. Stem imbricate-foliate, erect-
ascending when growing in dense mats, cherry-red, 5-
7 cm long; with weak central strand and thick-walled
epidermal cells; regularly pinnate-branched, branch-
es to 8 mm long, attenuate; in non-dense growth (on
tops of stumps, etc.), sympodial branching common,
with arcuate sympodial branches. Pseudoparaphyllia
broadly triangular to transversely ovate. Axillary hairs
4-6-celled, ca. 250 μm long, apical cell 40-50 × 11 μ m.
Stem leaves 1.8-2.6 × 1.0-1.4 mm, ovate, obtuse, pli-
cate when dry; margins incurved, entire to serrulate
above. Branch leaves 0.9-1.1 × 0.3-0.5 mm. Laminal
109
Hypnaceae and related pleurocarps with bi- or ecostate leaves in Altai
100 μm
Fig. 78. Pleurozium schreberi (Brid.) Mitt. (from Uedinennoye Lake 880 m Ignatov 0/977): 1 – upper laminal cells
of stem leaf; 2-5 – branch leaves; 6 – capsule; 7 – alar cells of stem leaf; 8 – habit; 9, 11 – pseudoparaphyllia; 10
– portion of stem with a branch; 12 – stem cross-section; 13-15 – perichaetial leaves; 16 – middle laminal cells of
stem leaf. Scale bars: 1 cm – for 8; 5 mm – for 6, 10; 1 mm – for 2-5; 100 μm – for 1, 7, 9, 11, 12, 16.
15
13
14
1
5
8
6
3
4
7910
11
12
2
16
1 cm
5 mm
1 mm
cells 50-105 × 8-10(6-7/2-4 ) μm. Alar cells subquad-
rate, forming a well-delimited, often colored group.
Dioicous. Sporophytes rather rare. Perichaetial
leaves 3.0 × 0.7 mm, erect, lanceolate, smooth, ecos-
tate. Seta 3 cm. Capsule ca. 2.5 mm long. Operculum
conic. Annulus none. Exostome teeth 0.55-0.60 mm
long, straight to ca. 2/3, distinctly narrowed in the
transition zone, entire; irregularly reticulate below;
in the middle densely papillose on few plates; in the
upper part with small low papillae arranged in in-
distinct longitudinal rows; dorsal trabeculae and
incrassate margin low-papillose; ventral trabeculae
smooth. Endostome 0.50 mm long, basal membrane
0.20 mm high; segments broadly perforate and of-
ten strongly slit; plates of the outer surface of seg-
ments jointed to the upper plates at apparent angle,
110 M. S. IGNATOV, H. ANDO & E. A. IGNATOVA
1 2
43
Fig. 79. Pleurozium schreberi (Brid.) Mitt. (from Uedinennoye Lake 880 m Ignatov 0/977): 1 – part of peristime
(400×); 2 – outer surface of exostome tooth in upper part (3400×); 3-4 – outer surface of exostome tooth in lower
part (3 – 6000×; 4 – 1100×).
resulting in peculiar transverse ridges. Cilia appen-
diculate. Spores 16-20 μm.
An easily known species by robust julaceous
plants with obtuse leaves and red stems.
Distribution: A very common moss in the Arctic,
boreal and hemiboreal regions of the Holarctic, P.
schreberi rarely occurs in temperate forests where it
grows mostly as an epixylic species. Outside the
Holarctic it is known in high mountains of Central
and South Americas. In Altai Pleurozium schreberi
occurs in nearly all the mesic forests and alpine com-
munities (up to the uppermost elevations studied).
It is amazingly absent in South-East Altai, on the
south-facing slope of Kuraiskij Mt. Range (the basin
of Chuya River). We failed to find it there in both
tundras and Larix (with Hylocomium + Rhytidi-
um) forests, though on the north-facing slope of that
range, in Bashkaus River Basin, it was found as a
very rare species.
Specimens examined: Bogoyash Creek, upper course 2200 m
(36/165); Bolshoye Istyube Creek 470 m (0/2089a); Boosh-
kon Creek, upper course 1900 m (Zolotukhin & al. 31.VII.1979);
Chiri Creek, 0.5 km upstream 450 m (17/21a); Kairu-Bazhi
Peak 2050 m (13/109); Karatumysh 2000 m (Zolotukhin & al.
30.VII.1978); Kayakkatuyarykskij Creek 1920 m (3/161a);
Kobiguayuk Creek 2220 m (0/360), 2300 m (0/652), 2908 m
(0/976); Kolyushta Peak 2030 m (Zolotukhin & al. 27.VII.1982);
Malaya Kokorya Creek 2300 m (36/168); Seminskij Pass 2000
m (173; Velikanov & al. 27.VIII.1961); Tamanel Peak 1800 m
(34/186); Turochak Creek, at mouth 970 m (Zolotukhin & al.
5.VI.1986); Uedinennoye Lake 880 m (0/977).
AKNOWLEDGEMENTS
We are very grateful to Dr. N. I. Zolotukhin and
L. V. Bardunov for making to our disposal their col-
lection from Altai and adjacent regions and also to
curators of H, FH, G, LE, NY, and S for loan of spec-
imes, including type collections important for this
study. Also we thank Dr. W. R. Buck for the revision
of English and other advises on the manuscri pt.
111
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